‚ivist— st—li—n— di €—leontologi— e ƒtr—tigr—fi— volume IIW noF Q ppF QPSEQSH xovem˜er PHIQ

ps‚ƒ„ ‚igy‚h yp pyƒƒsv g‰ƒ„y€ry‚sxei @ge‚xs†y‚eD €rygsheiAX

wshhvi wsygixi ƒievƒ p‚yw „ri xy‚„ri‚x €e‚e„i„r‰ƒ

s‚sxe eF uy‚i„ƒu‰ 8 ƒ vwex tF ‚erwe„

‚e™eivedX epril IPD PHIQY —™™eptedX tune PSD PHIQ

uey wordsX €ho™id—eD gystophorin—eD p—™hyosteos™lerosisD seE inusu—li per l9inter— f—migli—F sl nuovo genere del wio™ene ™ondivide

xu—l dimorphismD wio™eneD widdle ƒ—rm—ti—nD €—r—tethysD kr—ineF numerosi ™—r—tteri ™on diverse spe™ie ‚e™enti di gystophor— e wirounE

g—D ed offre l— prim— opportunit—Ádi studi—re il dimorfismo sessu—le delle

e˜str—™tF hespite — long history of pho™id studiesD no fossil oss— degli —rti e delle m—ndi˜ole nell— ƒottof—migli— gystophorin—eF sl

mem˜ers of the ƒu˜f—mily gystophorin—e h—ve ever ˜een des™ri˜edF dimorfismo sessu—le nelle oss— post™r—ni—li e nelle m—ndi˜ole eÁ m—gE

xew fossil m—teri—l from the widdle ƒ—rm—ti—n @IIFPEIPFQ w—A in the giormente evidente nei mem˜ri —ttu—li di gystophorin—e rispetto —gli

€—r—tethy—n f—sin of kr—ine —llows emended di—gnoses —nd redes™ripE —ltri r—ppresent—nti delle fo™he in senso strettoF v9—n—lisi dei tr—tti —n—E

tions to help ™l—rify phylogeneti™ rel—tionships within the p—mily €hoE tomi™i h— dimostr—to ™he entr—m˜e le nuove spe™ie sono piuÁprimitive e

™id—eF efter ™l—disti™ —nd morphologi™—l —n—lyses of the m—teri—lD — new meglio —d—tt—te per l— lo™omozione terrestre di qu—lsi—si —ltro r—ppreE

genus @€—™hypho™—A w—s ere™tedD with two new spe™ies of extin™t fossil sent—nte vivente di gystophorin—eF €—™hypho™— ukr—ini™—D piuÁpi™™ol—D eÁ

true se—ls @€—™hypho™— ukr—ini™— —nd €—™hypho™— ™h—pskiiAD ˜elonging meglio —d—tt—t— —ll— lo™omozione terrestre rispetto —ll— spe™ie piuÁgr—nE

to the ƒu˜f—mily gystophorin—eF „his new m—teri—l shows ex™eption—l de €F ™h—pskiiF v9—n—lisi filogeneti™— suggeris™e ™he le fo™he ™on IH in™iE

p—™hyosteos™leroti™ ˜onesD whi™h is un™ommon for the f—mily —s — sivi @€ho™in—eA sono piuÁprimitive di quelle ™on V @won—™hin—eAD e ™he i

wholeF „he new wio™ene genus sh—res numerous ™h—r—™ters with seE won—™hin—e sono piuÁprimitivi delle fo™he ™on T in™isivi @gystophoriE

ver—l ‚e™ent spe™ies of gystophor— —nd wiroung—D providing the first n—eAF uesti ritrov—menti indi™—no ™he l— ƒottof—migli— gystophorin—e

opportunity to study sexu—l dimorphism of lim˜ ˜ones —nd m—ndi˜les in™lude non solo elef—nti m—rini e fo™he d—l ™—ppu™™ioD m— —n™he le due

in the ƒu˜f—mily gystophorin—eF ƒexu—l dimorphism in post™r—ni—l nuove fo™he p—™hiosteos™leroti™he del ƒ—rm—ti—no wedioF

˜ones —nd m—ndi˜les in living mem˜ers of gystophorin—e is more o˜E

vious th—n in other represent—tives of true se—lsF ix—min—tion of —n—toE

mi™—l tr—its demonstr—ted th—t ˜oth new spe™ies —re more primitive —nd

˜etter —d—pted for terrestri—l lo™omotion th—n —ny living represent—tives sntrodu™tion

of gystophorin—eF „he sm—ller €—™hypho™— ukr—ini™— is more —d—pted

to terrestri—l lo™omotion th—n its l—rger rel—tiveD €F ™h—pskiiF €hylogeE „he study of fossil se—ls of the northern fl—™k ƒe—

th neti™ —n—lysis suggests th—t se—ls with IH in™isors @€ho™in—eA —re more ™entury with region @pigF IeA ˜eg—n in the mid IW

primitive th—n those with V @won—™hin—eAD —nd th—t won—™hin—e —re

work ˜y ii™hw—ld @IVSHD IVSQA —nd xordm—nn more primitive th—n se—ls with T in™isors @gystophorin—eAF „hese finE

@IVTHAD —nd w—s ™ontinued ˜y endrusov @IVWQAD elekE dings indi™—te th—t the ƒu˜f—mily gystophorin—e in™ludes not only

seev @IWPRAD ƒimiones™u @IWPSAD w—™—rovi™i —nd yes™u eleph—nt —nd hooded se—lsD ˜ut —lso the two new widdle ƒ—rm—ti—n

p—™hyosteos™leroti™ se—lsF @IWRIAD w™v—ren @IWTHAD uirpi™hnikov @IWTIAD uoretsky

@PHHIAD —nd uoretsky —nd qrigores™u @PHHPAF e diverse

‚i—ssuntoF xonost—nte l— lung— stori— dello studio dei fo™idiD

mix of true se—ls @€ho™id—eA h—s ˜een reported in these non er— m—i st—to des™ritto un mem˜ro dell— ƒottof—migli— gystophoE

studiesD ˜ut no mem˜ers of the yt—riid—eF rin—eF xuovo m—teri—le fossile d—l ƒ—rm—ti—no wedio @IIFPEIPFQ w—A del

hespite the —˜und—n™e —nd ˜ro—d distri˜ution of f—™ino dell— €—r—tetide dell9 ™r—in— permette or— di emend—re di—gnosi

e des™rizioni in modo d— —iut—re — ™hi—rire i r—pporti filogeneti™i —ll9inE €ho™id—eD m—ny pro˜lems persist in interpreting the

terno dell— f—migli— €ho™id—eF sn seguito —ll9—n—lisi morfologi™— e ™l—diE

system—ti™s of this groupD with the t—xonomi™ history

sti™— del m—teri—le eÁst—to istituito un nuovo genere @€—™hypho™—AD ™on

of gystophor— ˜eing espe™i—lly ™onfusing @uoretsky 8

due nuove spe™ie di fo™he in senso stretto estinte @€—™hypho™— ukr—ini™—

role™ PHHPAF qr—y @IVTTA —nd „rouess—rt @IVWUA divided e €—™hypho™— ™h—pskiiAD —pp—rtenenti —ll— ƒottof—migli— gystophorin—eF

uesto nuovo m—teri—le mostr— oss— p—™hiosteos™leroti™he e™™ezion—liD €ho™id—e into three su˜f—miliesX €ho™in—eD won—™hin—e

v—˜or—tory of ivolution—ry fiologyD hep—rtment of en—tomyD gollege of wedi™ineD row—rd niversity SPH ‡ ƒtF x‡D ‡—shingtonD hg PHHSW @ ƒeAF iEm—ilX ikoretIPQd—olF™omY sulm—nFr—hm—tdhow—rdFedu

QPT uoretsky sFeF 8 ‚—hm—t ƒFtF

pigF I E eA €—leogeogr—phi™ sket™hE

m—p of the i—stern €—r—tethys

for the dur—tion of the ƒ—rm—E

ti—n ƒe— @IPFQ ± WFQT w—A with

redu™ed s—linity —nd strong

endemism @m—p modified

—ftersv—nov et —lFD PHHUAF fA

qener—lized geogr—phi™—l m—p

of kr—ine showing the prinE

™ip—l fossil lo™—lities of the

genus €—™hypho™— from widE

dle wio™ene @ƒ—rm—ti—nA of

kr—ine mentioned in the textX

IA uhomutovo †ill—ge

@RUëIT9IIFR99x QVëHW9QTFI99iAF

PA qnylozu˜ovo

@RUëHT9STFQ99x QUëQI9TSFV99iAF

QA olot—y— f—lk—

@RUëPP9RTFW99 x QQëSV9PQFS99iFAF

RA heltok—menk—

@RUëRI9QRFP99x QQëSH9HTFV99iAF

SA „—r™h—nkut

@RSëPH9RTFW99x QPëPW9PQFS99iAF

TA zunl—r

@RSëIS9VHFI99x QTëPP9PUFH99iAF

UA qritsev

@RWëSV9HSFP99x PUëIH9HQ99iAF

—nd gystophorin—eF ‡hile there h—s never ˜een univerE in™isors to PGIY PA possession of —n infl—t—˜le n—s—l s—™Y QA

s—l —™™ept—n™e of this ™l—ssifi™—tionD it w—s followed ˜y nonEdifferenti—ted sh—pe @homodontyA of the post™—nine

m—ny su˜sequent workers in™luding ygnev @IWQSAD teethY RA sh—pe —nd dire™tion of m—stoid pro™essY —nd SA

qr—sse @IWSSAD ƒ™heffer@IWSVAD uing @IWTRA —nd gh—psE sh—pe of the m—xill—F ellen @IVVHAD followed ˜y ƒimpson

kii @IWURAF gh—r—™ters supporting the existen™e of three @IWRSAD ™re—ted —n —ddition—l su˜f—milyD vo˜odontin—eD

su˜f—milies m—inly derive from —n—lyses of ™r—ni—lD denE for southern se—ls th—t gh—pskii —nd previous workers

t—l —nd pel—ge morphologyF sn p—rti™ul—rD — ™omprehenE reg—rded —s — tri˜e within won—™hin—eF

sive —n—lysis of supr—generi™ system—ti™s is found in the w—jor divergen™es from the threeEsu˜f—mily

studies of gh—pskii @IWSSD IWTID IWUID IWURAF re deE s™heme ˜eg—n with the work of uing @IWTTA —nd furns

s™ri˜ed di—gnosti™ ™r—ni—l ™h—r—™ters @num˜er of in™iE —nd p—y @IWUHAF gh—pskii @IWURA —lso proved th—t uing9s

sorsD sh—pe of m—xill—D form of —nterior p—l—t—l for—min—D @IWTTA hypothesis of re—ssigning the genus gystophor—

—nd some othersA supporting the sep—r—tion of true se—ls from the ƒu˜f—mily gystophorin—e into the ƒu˜f—mily

into three su˜f—miliesD whi™h he in turn divided into €ho™in—e —nd the genus wiroung— into the ƒu˜f—mily

tri˜es —nd su˜Etri˜esF won—™hin—e is unten—˜leF wuizon @IWVPA —™™epted the

„hereforeD se—ls with six in™isors ˜elong to the system—ti™s of uingD —nd returned to dismissing the ƒu˜E

ƒu˜f—mily gystophorin—e @qill IVTTAD ™omprising the f—mily gystophorin—e —s — wholeF „he ™orre™tness of

gener— gystophor— xilssonD IVPH @hooded se—lA —nd gh—pskii9s ™on™ept w—s ™orro˜or—ted ˜y ‚o˜inette

wiroung— qr—yD IVPU @eleph—nt se—l or se— eleph—ntAF —nd ƒt—ins @IWUHA —nd €olly @PHHVA in their™omp—r—tive

„his ™l—ssifi™—tion is ˜—sed on sever—l syn—pomorphies studies of the pinniped ™—l™—neusD emph—sizing th—t it is

˜etween gystophor— —nd wiroung—X IA redu™tion of the in—dmissi˜le to t—xonomi™—lly sep—r—te the hooded se—l

widdle wio™ene se—ls from the xorthern €—r—tethys QPU

—nd the se— eleph—ntF v—terD this point of view w—s —lso teri—l @—s for tod—yD only IH known fossil se—l skulls h—ve

supported ˜y en˜inder @IWVHXUTAX ––FFF —n—lyti™—l meE ever ˜een found —nd des™ri˜edA is th—t the surviv—l r—te

thods of ™hromosome investig—tions —™tu—lly do not perE of fossil se—l skulls is extremely low due to the p—perE

mit the sep—r—tion of gener— gystophor— —nd wiroung—D thin thi™kness of skull ˜onesF e simil—r™ondition —pE

—nd this ™ontr—sts with the ™on™ept of their sep—r—te pe—rs in ot—rid—eD ˜ut fossil ot—rid—e skull ˜ones —re

t—xonomi™ st—tus —nd of in™lusion of gystophor— in mu™h denser —nd h—ve — higher surviv—l r—teF

€ho™in—e99F roweverD re™ent mole™ul—r —nd k—ryologi™—l hue to the gre—t r—rity —nd usu—lly uns—tisf—™tory

studies do not support this point of view —nd pl—™e preserv—tion of post™r—ni—lD —nd espe™i—lly ™r—ni—lD fossil

gystophor— into the ƒu˜f—mily €ho™in—e —nd wiroung— rem—ins of se—lsD ™omp—red to terrestri—l ™—rnivoresD this

into the ƒu˜f—mily won—™hin—e —s sep—r—te ™l—des @fiE rem—ins one of the le—st investig—ted —nd most ™ompliE

È nind—Eimonds 8 ‚ussell IWWTY ern—sson et —lF PHHTY ™—ted groups of l—rge m—mm—lsF „his study fo™used on

pulton 8 ƒtro˜e™k PHIHAF iven mole™ul—r ˜iologists des™ri˜ing previously unknown extin™t represent—tives

™on™lude the need for— reEev—lu—tion of pinniped t—xoE of gystophorin—e —nd ™l—rifying their rel—tionships with

nomy @rigdon etF —lF PHHUAF yurmorphologi™—ld—t— otherse—ls of the p—mily €ho™id—e —nd its foursu˜E

indi™—tes th—t the new fossil se—ls from the widdle wioE f—milies @€ho™in—eD won—™hin—eD gystophorin—eD —nd

™ene @€—™hypho™—A h—ve — ™om˜in—tion of ™ystophorine hevinopho™in—eAF

—nd miroungini ™h—r—™tersD reEopening the dis™ussion emong xeogene m—rine m—mm—lsD the rem—ins

on™e —g—in —˜out the su˜f—mili—l rel—tionships of se—lsF of pho™ine se—ls @in ™omp—rison with mon—™hines —nd

uing @IWTRAD furns —nd p—y @IWUHAD g—rr —nd €erE ™ystophorinesA —re rel—tively numerous in the widdle

È ry @IWWRAD —nd l—ter ern—son et —lF @PHHTA —ssigned gysE ƒ—rm—ti—nEw—eoti—n deposits of the gentr—l —nd i—stern

tophor— to the „ri˜e gystophoriniD while w™uenn— —nd €—r—tethys @pigF IAF ƒu™h rem—ins —re found in the kE

fell @IWWUA in™luded gystophor— in the „ri˜e €ho™iniF r—ineD wold—vi—D ‚om—ni—D u—z—khst—nD ƒlov—ki—D eusE

iven tod—yD the ex—™t t—xonomi™ rel—tionships of se—ls tri—D rung—ryD —nd even in „urkey @ii™hw—ld IVSHY

within €ho™id—e —re still ™ontroversi—l @‡yss IWWRY uorE xordm—nn IVTHY elekseev IWPRY ƒimiones™u IWPSY

etsky 8 qrigores™u PHHPY uoretsky 8 role™ PHHPAF w™v—ren IWTHY qrigores™u IWUUY qrigores™u et —lF

†—rious s™ientists either sep—r—te pho™ids intoX only IWVTY uoretsky IWVTD IWVU—D ˜D PHHIY uoretsky 8 qriE

one su˜f—milyD €ho™in—e @‡yss IWVVY w™uenn— 8 fell gores™u PHHPAF sn spite of the —˜und—n™e —nd ˜ro—d disE

IWWUAY two su˜f—miliesD €ho™in—e —nd won—™hin—e tri˜ution of true se—lsD m—ny pro˜lems persist in interE

@furns 8 p—y IWUHY wuizon IWVPD IWWPY uing IWVQD preting the system—ti™s of these —nim—ls @uoretsky

IWVWY ‡yss IWWRY €erry et —lF IWWSY finind—Eimonds IWVU—AF emong the three ––tr—dition—l99 su˜f—milies of

8 ‚ussell IWWTY fert— 8 ƒumi™h IWWWAY or do not sep—E €ho™id—e @€ho™in—eD won—™hin—eD gystophorin—eA —nd

r—te true se—ls into su˜f—milies —t —ll @ƒokolov IWUWY the more re™ently des™ri˜ed su˜f—mily hevinopho™in—e

‡ozen™r—ft IWVWAF „hereforeD this study will ™ontinue @uoretsky 8 role™ PHHPAD fossil rem—ins of gystophoriE

to follow the ––™l—ssi™—l99 ™l—ssifi™—tion of ƒ™heffer—nd n—e h—ve never ˜een foundD illustr—ted or des™ri˜edF

gh—pskii ˜y pl—™ing gystophor— into the ƒu˜f—mily gysE „he extensive investig—tions of fossil se—ls from the

tophorin—e ± the sixEin™isor se—lsF widdle wio™ene deposits of kr—ine now —llow — more

fe™—use of the p—u™ity of ™r—ni—l rem—insD the —™™ur—te di—gnosis of the ƒu˜f—mily gystophorin—eD —nd

study of fossil se—ls h—s ˜een ˜—sed m—inly on the use ™omp—risons ˜etween this t—xon —nd other gener— of the

of post™r—ni—l ™h—r—™tersD spe™ifi™—lly the morphology of p—mily €ho™id—e ™—n help determine the phylogeneti™

individu—l disso™i—ted ˜ones @m—inly the femurorhuE rel—tions —mong the su˜f—miliesF

merusAF snterpret—tion of post™r—ni—l elements is —ided „his study will improve the ™l—ssifi™—tion —nd

˜y —n—lysis of spe™ifi™ e™ologi™—l ni™hes th—t —re reE knowledge of the gener—l morphology of true se—lsD

fle™ted in ˜ones of the post™r—ni—l skeleton —nd m—ndiE with spe™i—l emph—sis on previously unknown fossil reE

˜le of ‚e™ent se—ls @uoretsky PHHIAF „his h—s —llowed present—tives of the ƒu˜f—mily gystophorin—e from the

sep—r—tion of modern pho™ines into morphologi™—l widdle ƒ—rm—ti—n @IPFQEIIFP w—A of the i—stern €—r—E

groups ˜—sed on ™h—r—™ters from the most ™ommon disE tethysD not—˜ly in the northern fl—™k ƒe— littor—l region

so™i—ted elements @more th—n IHHH ˜onesD m—inly m—ndiE of the kr—ineF „—xonomi™ ™h—r—™ters used for the ™l—sE

of €ho™id—e @uoretsky PHHIY uoretsky 8 qriE ˜leD humerusD —nd femurAF sn —dditionD other pu˜li™—E sifi™—tion

gores™u PHHPY uoretsky 8 ‚—y PHHVA will ˜e —n—lyzed in tions show —sso™i—ted p—rts of se—l skeletons @wuizon

order to determine whi™h su˜f—mily our two new speE IWVIY uoretsky PHHIY gozzoul PHHIY uoretsky 8 qriE

™ies ˜elong in —nd the rel—tionships they h—ve with other gores™u PHHPY uoretsky 8 ‚—y PHHVA th—t —lso ™—n ˜e

true se—lsF „husD the new fossil eviden™e presented in used —s — found—tion of —lph— ™l—ssifi™—tionF „his inforE

this study —lso ™—n ˜e used to determine the rel—tionE m—tion w—s the ˜—sis to —sso™i—te individu—l ˜ones from

ships within the ƒu˜f—mily gystophorin—eD —nd proE the widdle ƒ—rm—ti—n of kr—ine into two t—xonomi™—l

vides — unique opportunity to study sexu—l dimorphism unitsF enother hurdle for finding ™r—ni—l fossil se—l m—E

QPV uoretsky sFeF 8 ‚—hm—t ƒFtF

—t some lo™—litiesF „hese —sso™i—tions give the potenti—l of the lim˜ ˜ones —nd m—ndi˜les in living mem˜ers of

this su˜f—milyF for two ™rossE™he™king systems @one through l—nd reE

™ords vi— esi—D —nd the other through m—rine re™ords vi—

e˜˜revi—tions

the xorth etl—nti™A of ˜io™hronologi™—l ™orrel—tion ˜eE

ƒpe™imens from the following institutions —nd dep—rtments

tween iur—si— —nd xorth emeri™— @uoretsky et —lF h—ve ˜een ex—mined forthis m—nus™riptX

PHIPAF ewxrD emeri™—n wuseum of x—tur—l ristoryD xew ‰orkD

ƒeF „he geogr—phi™ r—nges of some modern pinniped

xwxr E€D x—tion—l wuseum of x—tur—l ristory —t the x—E

spe™ies —re very l—rge —nd often extend through sever—l

tion—l e™—demy of ƒ™ien™e of kr—ineD uievD kr—ineF

zoogeogr—phi™—l regionsF vikewiseD fossil spe™ies —re

ƒxwD x—tion—l wuseum of x—tur—l ristoryD ƒmithsoni—n

widespre—dF „rue se—ls @€ho™id—eA —re of p—rti™ul—r zooE snstitutionD ‡—shingtonD hFgFD ƒeF

geogr—phi™ interest —s their fossils —re very numerous in sxD oologi™—l snstitute of the ‚ussi—n e™—demy of ƒ™ien™esD

ƒtF €eters˜urgD ‚ussi—F wio™ene ™o—st—lEm—rine f—un—s of the northern fl—™k

sn referen™e to ˜onesX

ƒe— littor—l regionF sn the former ƒoviet nionD rem—ins

vF ± leftY ‚F E rightD mF ± mus™leF

of these —nim—ls h—ve ˜een found in the „r—ns™—u™—sus

—nd u—z—khst—nD with the most numerous finds from

the northern littor—l region of the fl—™k ƒe— in the kE

xorthern fl—™k ƒe— m—rine m—mm—l lo™—lities

r—ine —nd wold—vi— from the widdle ƒ—rm—ti—n @IPFQ ±

IIFP w—AF sn ‡estern —nd gentr—l iuropeD rem—ins of

„he wediterr—ne—nD fl—™kD g—spi—n —nd er—l ƒe—s

fossil se—ls —re regul—rly found in pr—n™e @pri—nt IWRPD

—re remn—nts of the —n™ient „ethys se—w—y whi™hD duE

IWRUY qins˜urg 8 t—nvier IWUSD IWWWAD „he xetherl—nds ring mu™h of wio™ene time @PQ to SFP w—AD extended

@ƒpringhorn IWUVY uoretsky 8 €eters PHHVY uoretsky et

™ontinuously from ‡estern iurope into gentr—l esi—

—lF PHIPAD felgium @uoretsky 8 ‚—y PHHVAD henm—rk

@pigF IeAF „he northern shore of this se—w—y extended

@uoretskyD in pressAD eustri— @—pfe IWQUAD rung—ry

—™ross the —re— th—t is now southern kr—ineF ƒediments

@uoretsky PHHQAD ‚om—ni— @qrigores™u IWUUY uoretsky deposited inD —nd on the shores ofD this —n™ient se—w—y

8 qrigores™u PHHPAD gze™h ‚epu˜li™ @role™ et —lF IWVUY —re ri™h in the rem—ins of fossil m—mm—lsF xe—r the

role™ 8 ƒ—˜ol IWWTY role™ et —lF IWWUAD —nd ƒlov—ki— present northern shore of the fl—™k ƒe— —re thi™k se™E

@uoretsky 8 role™ PHHPAF ssol—ted finds —re known tions of predomin—ntly m—rine str—t—D with sever—l loE

from „urkey @uoretskyD unpu˜lished d—t—AD st—ly @„—v—E ™—lities @„—r™h—nkutD uer™hD ydess—A h—ving —n —˜unE

ni IWRPA —nd w—lt— @fi—nu™™i etF —lF PHIIAF possil wioE d—n™e of fossil m—rine m—mm—l rem—insD in™luding pinE

™ene se—ls —re —lso known in the e—stern nited ƒt—tes nipeds @se—lsA —nd ™et—™e—ns @wh—lesD porpoisesAD —s well

@uoretsky —nd ‚—yD PHHVAD with the most primitive —s some ™ontinent—l deposits with l—nd m—mm—l rem—ins

˜eing veptopho™— lenis „rueD IWHTD from the widdle @€idopli™hko IWSTY hu˜rovo 8 u—pelist IWUWY uoretsky

wio™ene of w—ryl—nd —nd †irgini— @‚—y IWUTY uoretsky IWVTD IWVUAF

PHHIY uoretsky et —lF PHIPA —nd re™ently dis™overed in p—rther to the northD deposits —re predomin—ntly

xorth efri™— in vi˜y— @uoretsky —nd homningD in ™ontinent—l with rem—ins of l—nd m—mm—ls preserved —t

pressAF m—ny lo™—lities @ƒev—stopolD †elikomi™h—ilovk—D feriE

w—ny m—rine —nd ™ontinent—l xeogene deposits sl—vD qre˜enikiD eugustovk—D gu˜—nk—AF „he sever—l loE

h—ve not ˜een thoroughly studiedF €—rti™ul—rly insuffiE ™—lities @qri™evD olot—y— f—lk—D heltok—menk—D „y—E

™ient —re studies of xeogene m—rine deposits of the gink—Y pigF IfA where l—nd m—mm—ls —nd m—rine m—mE

kr—ineD espe™i—lly their str—tigr—phi™ ™orrel—tion with m—ls —re —sso™i—ted @uoretsky PHHIA —re espe™i—lly imE

™ontinent—l deposits of —dj—™ent —nd dist—nt regionsF port—ntF

„his study will ™on™entr—te upon wio™ene @PQFV ± SFP et ™ert—in interv—ls during the xeogeneD the fossil

w—A m—rine deposits in southwestern kr—ineD whi™h re™ord shows th—t the l—nd m—mm—li—n f—un— of the

—re the most extensively representedD ˜ut le—st ™h—r—™E northern ™ontinents w—s quite ™osmopolit—nD indi™—ting

terized geologi™—llyF „he thi™k m—rine l—yers exposed in periods when it w—s possi˜le for l—nd m—mm—ls to disE

this region —reD —lmost —s — ruleD not very ri™h in fossil perse freely ˜etween iurope —nd esi— —s well —s to

—nim—l rem—ins other th—n mollusksF sn simil—rE—ged xorth emeri™— vi— feringi— @fernor etF —lF IWWHAF x—tuE

™ontinent—l deposits of the kr—ineD some l—rge wioE r—llyD the dispers—l p—ttern for m—rine m—mm—ls is very

™ene hipp—rion f—un—s h—ve ˜een foundD suggesting th—t differentD —s fossil se—ls —nd wh—les of the „ethys ™ould

southwestern kr—ine is —n import—nt lo™—tion for p—E r—nge —™ross the xorth etl—nti™ to e—stern xorth emeE

leozoologi™—l investig—tionsF ri™—n ™o—st—l w—ters @uoretsky 8 f—rnes PHHTY uoretsky

ƒin™e elekseev @IWPRAD — signifi™—nt fossil se—l 8 ‚—y PHHVY uoretsky et —lF PHIPAF possil se—l t—x— found

™olle™tion from i—stern €—r—tethys h—s —™™umul—ted in in €—r—tethy—n lo™—lities —™ross iurope —lso h—ve ˜een

dis™overed in deposits —long the e—stern se—˜o—rd of uiev9s x—tion—l wuseum of x—tur—l ristory —t the x—E

xorth emeri™—D with terrestri—l m—mm—li—n —sso™i—tion tion—l e™—demy of ƒ™ien™eF „hese fossils h—ve ˜een reE

widdle wio™ene se—ls from the xorthern €—r—tethys QPW

€ho™— spF @€idopli™hko IWSTY hu˜rovo 8 ferred to —s Pachyphoca

Characters

u—pelist IWUWAF roweverD this ™olle™tion of very p—E ukrainica chapskii

™hyosteos™leroti™ post™r—ni—l elements must ˜e revised Maximal length 118.5 121.0

—nd ––redes™ri˜ed99 in order to ˜ro—den gener—l knowlE Anteroposterior diameter of cavitas glenoidalis 17.5; 18.0 19.0 edge —nd underst—nd the spe™ifi™ distri˜ution of ™ystoE Transverse diameter of cavitas glenoidalis 21.0; 21.0 30.0

Maximal thickness of sapula in its spine 21.0; 19.0 24.5

phorines —nd their rel—tionship with other su˜f—miliesF Width of cervix 21.5; 24.0 26.5

„his revision will yield — mu™h more pre™ise —nd deE Transverse width of body 60.0 75.5 t—iled di—gnosis of the m—teri—l —nd deline—te the distriE

„—˜F I E we—surements @mmA of s™—pul—eF ˜ution of this t—xonF „he inform—tion presented ˜elow

on geogr—phi™—l lo™—tionsD their geologi™—l —geD ™olle™E

pigF P E ‚ostr—l p—rt of ‚e™ent se—l

m—xill—e in eA hors—l views

@modified —ftergh—pskiiD

IWURAF v—˜i—l views of gystoE

phor— ™rist—t— m—ndi˜lesX fA

m—le @ ƒxw PTWIQHY reE

versedA —nd gA fem—le

@ ƒxw IVVWSTAF

e˜˜revi—tionsX intD n—s—l proE

™ess of interm—xill—Y m—xD m—xill—F

QQH uoretsky sFeF 8 ‚—hm—t ƒFtF

torsD —nd institutions where m—teri—l is stored w—s ™omE —nd two new spe™ies of the ƒu˜f—mily gystophorin—eD

piled using pu˜lished d—t— @€idopli™hko IWSTY hu˜rovo —ll housed in the hep—rtment of €—leontology of the

xwxr E€F 8 u—pelist IWUWAF

i—rlierD uoretsky @PHHIA showed th—t there —re

sever—l lo™—lities @qri™evD ghomutovoD olot—y— f—lk—D

heltok—menk—D —nd „y—gink—A in kr—ine where se—l ƒystem—ti™ €—leontology

˜ones with pronoun™ed p—™hyosteos™lerosis —re espeE

™i—lly noteworthy @pigF IfAF „hese —nd others —re —s

ƒuperf—mily €ho™oide— ƒmirnovD IWHV

followsX

p—mily €ho™id—e qr—yD IVPS

heltok—menk—X hnepropetrovsk regionD eposE

ƒu˜f—mily gystophorin—e qr—yD IVTT tolovo distri™tD heltok—menk— vill—ge on eltenk—y—

‚iverD limestone qu—rry VEIH m deepY widdle ƒ—rm—ti—n

„ype qenusX gystophor— xilssonD IVPHY present distri˜utionX

@fess—r—˜i—nY IPFQ ± IIFP w—AY expeditions of €idopli™hE

er™ti™ —nd su˜—r™ti™ xorth etl—nti™ o™e—nsY ™ir™umpol—r in su˜—nt—r™E

ko IWQVD IWRHY fezuglui IWSQESRY —nd ƒemenov PHHIY ti™ regionF

™olle™tion of xwxr E€F ytherm—mm—li—n rem—insD histri˜utionX widdle wio™ene @ƒ—rm—ti—nA in i—stern €—r—E

tethysY ‚e™ent in er™ti™ —nd ent—r™ti™ y™e—nsF ˜esides pinnipedsD su™h —s w—™h—irodusD w—stodonD

imended hi—gnosisX v—rge se—ls @length up to S mA with six

e™hti—ri— —nd getotheium were found in this qu—rry

in™isors @saPGIY in ™ontr—st to won—™hin—e with V in™isors —nd €ho™in—e

in the IWRHsF ƒome ˜ones of the m—rine m—mm—ls @se—ls

with IH in™isorsAY p—ro™™ipit—l pro™ess of skull poorly developed @in

—nd wh—lesA —re very mu™h rolled —nd —˜r—dedD whi™h ™ontr—st to hevinopho™in—eAY —nteroposterior length of —uditory ˜ull—

less th—n dist—n™e ˜etween ˜ull—e @in ™ontr—st to €ho™in—e —nd hevinoE me—ns th—t they were deposited in sh—llow w—ter ne—r

pho™in—eAY infr—or˜it—l pro™ess presentY interor˜it—l sp—™e wideY interE the shore @shelf zoneAF

or˜it—l width less th—n QH7D ˜ut equ—l to or gre—ter th—n PS7 of

uhomutovo †ill—geX honetsk regionD xovo—zov

m—stoid width @—s in hevinopho™in—eAY —nterior p—l—t—l for—min— ov—l

distri™tY limestone qu—rry TEU m deepY pper wio™ene

@—s in hevinopho™in—eA —nd sh—llowY preor˜it—l p—rt of m—xill— with

@€onti—nAY ™olle™tion of IWSWY xwxr E€F n—rrow ™on™—vity @simil—r to vo˜odontiniY in ™ontr—st to won—™hin—eD

€ho™in—eD —nd hevinopho™in—eAF pper se™ond in™isors tend to enl—rge „—r™h—nkutX grime— regionD ghernomorsk disE

r—ther th—n first in™isorsF tri™tD „—r™h—nkut €eninsul—D U km southEe—st of vill—ge

w—ndi˜ul—r™hin prominen™e—˜sentY —lveoli of pR ˜iggerth—n

of ylenevk—Y widdle ƒ—rm—ti—nY ™olle™tions of entoniuk

—lveoli of mIY ™oronoid pro™ess very n—rrow —nd turned ™—ud—llyD espeE

IWUSEIWVTY —nd uoretsky IWVQEIWVTY sxD xwxr E€F ™i—lly in m—leY ™ondyloid pro™ess not well m—rkedY symphysis re—™hes

zunl—rX grime—D uer™h €eninsul—Y widdle ƒ—rE posterior —lveolus of pIY m—ndi˜ul—r not™h very n—rrowY retrom—ndi˜uE

l—rsp—™e elong—ted @~QFS ™m in fem—les —nd ~ Q ™m in m—lesAF m—ti—nY ™olle™tion IWSPY xwxr E€F

widdle of intern—l ™rest of humer—l tro™hle— rises w—veElike over

olot—y— f—lk—X gherson regionD xovovorontsoE

™oronoid foss—Y widths of dist—l —nd proxim—l epiphyses —lmost equ—lF

vo distri™tD right ˜—nk of the hnieper ‚iverD south of

wedi—l —nd l—ter—l femor—l ™ondyles —lmost equ—l in sizeY lesser

™ity of xikopolY limestone qu—rry north of vill—geD tro™h—nter present in m—lesY minimum width of femor—l sh—ft IFREIFW

upperl—yerDU m deep D widdle ƒ—rm—ti—nY ™olle™tion of times width of proxim—l epiphysisF

sn™luded qener—X gystophor— xilssonD IVPHY wiroung— qr—yD fezuglui IWSPY xwxr E€F

IVPUF

qnylozu˜ovX honetsk regionD w—riupol distri™tD

left ˜—nk of u—lmius ‚iverY vill—ge of fo˜rinetsY limeE gomp—risonsF „he interor˜it—l portion of the

stone qu—rryD widdle ƒ—rm—ti—nY ™olle™tion of IWQTY front—l ˜one in gystophorin—e @—s in hevinopho™in—eAD

xwxr E€F ˜ut in ™ontr—st to €ho™in—eD is widened @—pomorphi™ or

qritsevX ghmelnitsky regionD ƒhepetovk— distri™tD derived ™onditionAD even to — gre—ter degree th—n in

Q km west of vill—ge of qritsevY k—rst deposits in limeE won—™hin—eF yn the —nterior p—rt of the front—l ˜oneD

stone qu—rry on right ˜—nk of ghomor— ‚iverY widdle the origin of the tempor—l ™rest is presentD in ™ontr—st to

ƒ—rm—ti—nY ™olle™tion of ƒemenov IWVQEIWVSY the other three su˜f—miliesF „he p—rt of the m—xill— @pigF

xwxr E€F PeA situ—ted ˜etween the n—s—l me—tus —nd or˜its h—s —

xovovorontsovoX gherson regionD xovovorontE n—rrow ™on™—vityD —s in hevinopho™in—eD —nd ™ontr—sts

sovsky distri™tY ™l—yY widdle wio™eneY ™olle™tion of feE with €ho™in—e @with — short ™onvexityA —nd won—™hin—e

zuglui IWSPY xwxr E€F @with — long ™on™—vityAF sn ™ontr—st to the other three

xikol—evX xikol—ev regionD vi™inity of ™ity of xiE su˜f—miliesD —n interv—l is —˜sent ˜etween the extern—l

kol—evY i—rly ƒ—rm—ti—nY ™olle™tion of IWQSY xwxr E€F —uditory me—tus —nd the —nterior ™rest of the m—ndi˜uE

l—r foss—F „he l—™rim—l ˜one h—s —n —nteroEor˜it—l proE

„he m—teri—l w—s ™olle™ted —nd gr—du—lly —™™umuE ™ess strongly protruding to — gre—ter degree th—n in

l—ted during m—ny ye—rs @IWQP ± PHHQA of ex™—v—tions won—™hin—eF „he ™onvexity of the m—stoid is not roE

™—rried out ˜y expeditions of the hep—rtment of €—leonE ˜ustly expressed —nd is dire™ted ™r—nioE™—ud—llyF „he

for—men ov—le is ventr—lly ™overed ˜y the presphenoid tology of the xwxr E€F „he ™olle™tion in™ludes SV

˜oneD —nd l—ter—lly ˜y the postglenoid pro™essF „he individu—l post™r—ni—l ˜ones representing the new genus

widdle wio™ene se—ls from the xorthern €—r—tethys QQI

pigF Q E „ip of the —rrow points to the

middle of the humer—l troE

™hle—r™restYeA in €ho™in—eD

—t the level of the ™oronoid

foss—Y fA in won—™hin—eD

—r™hElike elev—ted over the

™oronoid foss—Y gA in gystoE

phorin—eD w—veElike r—ised

overthe ™oronoidfoss—F

—nteroEinferior pro™ess of the jug—l ˜one is lo™—ted —lE his™ussionF ythert—x— h—ve ˜een suggested to ˜e

most —t the level of the middle of the or˜itD in ™ontr—st ™losely rel—ted to mem˜ers of gystophorin—eF —pfe

to the other su˜f—miliesF „he —nterior p—l—t—l for—min— @IWQUA proposed th—t the represent—tives of the genus

—re ov—l —s in hevinopho™in—e —nd tend to dis—ppe—r —s wiopho™— —re undou˜tedly —n™estr—l to gystophorin—eF

in won—™hin—eF ‡hile ƒimpson @IWRSAD „henius @IWSHD IWSPAD uing

„he upper se™ond in™isorsD in ™ontr—st to the other @IWTRAD —nd w™uenn— —nd fell @IWWUA —ssigned wiophoE

su˜f—miliesD —re mu™h ˜igger th—n the first onesD —nd ™— vetust— to the ƒu˜f—mily won—™hin—eD otherinvestiE

—lmost re—™h the size of the ™—ninesF „he teeth —re very g—tors @‚—y IWUUY wuizon IWVPY ƒ—v—ge 8 ‚ussell IWVQA

sm—ll in ™omp—rison to the size of the m—ndi˜leD pR is

did not mention this t—xon —t —ll in theirreviewsof the

˜igger th—n mID ˜ut the l—rgest tooth is pQY €PD pP —re

„erti—ry se—ls of iuropeF „henius @IWSHD IWSPAD without

singleErootedY pP ± mI h—ve fused rootsD —nd the —lveoli

—ny expl—n—tionD tr—nsferred wiopho™— vetust— to —nE

—re sh—llowF

othergenusD €ristipho™—F v—terD role™ et —lF @IWVUD

sn gystophorin—eD the ˜ody of the m—ndi˜leD —nd

IWWTA supported this opinion with — propos—l to re—ssign

espe™i—lly the r—musD —re very uniqueD ™omp—red to the

genus wiopho™— to the „ri˜e gystophoriniF roweverD

othersu˜f—miliesF „he m—ndi˜ul—r˜ody is verythi™kD

the distin™tive morphologi™—l m—ndi˜ul—r ™h—r—™ters

˜ut not highF „he r—mus of the m—ndi˜le is —lso very

@uoretsky et —lF in pressAD espe™i—lly the simil—r degree thi™kD wideD —nd high in ™ontr—st to the other su˜f—miliesD

of the ™ondyloid —ngle @IPëEISëAD shows th—t wiopho™—

—s is the —˜sen™e of the m—ndi˜ul—r™hin prominen™eF

vetust— ™le—rly ˜elongs to the ƒu˜f—mily won—™hin—eF

„he ™ondyloid pro™ess is espe™i—lly shortD n—rrowD —nd

not pronoun™edF elthough we ˜elieve th—t gystophorE

in—e —nd wiroung— ˜elong to the s—me t—xonD some

ƒexu—l himorphism in gystophor— ™rist—t— e™omorphologi™—l ™h—r—™teristi™s do differD su™h —s the

gystophorine ™oronoid pro™ess of the m—ndi˜le ˜eing

sn prior studies of true se—lsD sexu—l dimorphism very sleek —nd turned ™—ud—llyD the presen™e of — very

w—s only ex—mined on ™r—ni—l m—teri—l of ‚e™ent €hoE n—rrow m—ndi˜ul—r not™hD —nd —n elong—ted retrom—nE

™in—e @ygnev IWQSY gh—pskii IWSPD IWTUY uhuzin IWTUAF di˜ul—rsp—™eD ™omp—redto those found in wiroung—F

roweverD fossil rem—ins of se—ls usu—lly ™onsist of isoE

„he middle of the humer—l ™rest of the tro™hle—

l—ted lim˜ ˜onesD m—inly the femur—nd humerusF„he

@pigF QA in gystophorin—e is w—veElike —nd elev—ted over

l—™k of fossil ™r—ni—l m—teri—l m—kes the use of post™r—E the ™oronoid foss—D in ™ontr—st to th—t in €ho™in—eD

ni—l elements even more import—nt in the proper ™l—ssiE whi™h is —t the level of the ™oronoid foss—D or in woE

fi™—tion of spe™imens —nd in ex—mining the sexu—l v—ri—E n—™hin—eD where it is —r™hElike —nd ™on™—vely r—ised over

˜ility @dimorphismA of v—rious ˜onesF ƒtudies determinE the ™oronoid foss—F

ing ontogeneti™ —nd sexu—l v—ri—tion ˜—sed on post™r—E „he femor—l ™ondyles —re —lmost equ—l in size @˜ut

ni—l elements of modern —nd fossil se—ls were done ˜y less so th—n in won—™hin—eAD in ™ontr—st to €ho™in—eF

q—djiev IWVPD uoretsky IWVU—D †—n free —nd ird˜rink „he minimum width of the di—physis is —lmost twi™e

IWVU —nd uoretsky PHHIF —s gre—t —s the width of the proxim—l epiphysisF

QQP uoretsky sFeF 8 ‚—hm—t ƒFtF

Pachyphoca Cystophora cristata Characters ukrainica chapskii female male n M OR n M OR n M OR Absolute length 2 85.0; 89.0 - 4 148.2 143.0-153.3 3 153.1 152.0-154.0 Length of deltoid crest 2 55.0; 56.0 - 4 81.0 78.0-83.9 3 83.3 82.6-83.4 Height of head 4 20.8 19.5-23.0 - 4 30.4 28.9-31.5 3 30.7 28.7-34.5 Height of trochlea 3 15.0 14.0-16.0 21.0 4 29.5 28.0-31.0 3 27.8 27.7-28.0 Width of head 4 23.3 22.0-25.0 - 4 34.5 34.0-35.1 3 35.9 34.6-38.0 Width of deltoid crest 2 19.5; 20.0 - 4 36.8 31.0-41.5 3 45.5 44.1-50.0 Width of distal epiphysis 2 33.0; 34.5 - 4 57.3 55.0-60.0 3 57.2 56.7-63.0 Width of proximal epiphysis 2 29.0; 32.0 - 4 53.8 49.5-58.0 3 56.0 53.5-60.0 Width of trochlea below 3 18.0 17.0-19.0 - 4 31.3 30.6-32.0 3 33.1 30.6-38.0 Width of trochlea, frontal view 3 21.3 20.0-23.0 49.5 4 31.7 28.0-34.0 3 35.9 35.0-37.7 Transverse width of diaphysis 4 18.3 16.5-21.5 27.0 4 23.6 23.2-24.0 3 23.9 22.2-27.0 Thickness of proximal epiphysis 2 29.0; 35.0 - 4 49.1 44.7-53.5 3 53.9 53.7-68.0 Thickness of medial condyle 3 16.3 15.5-17.0 23.0 4 25.0 24.0-26.1 3 25.7 24.1-27.0 Thickness of lateral condyle 3 15.2 15.0-5.5 20.0 4 28.2 26.4-30.0 3 27.8 27.0-31.0

Diameter of diaphysis with deltoid crest 2 31.5; 33.0 - 4 53.8 53.0-55.0 3 58.4 53.2-66.0

„—˜F P E we—surements @mmA of humeriF

ƒexu—l dimorphism in the post™r—ni—l ˜ones —nd in m—les @see det—iled expl—n—tion in uoretsky et —lF in

m—ndi˜les of the ƒu˜f—mily gystophorin—e @pigF RY „—˜s pressAF

PD TAD espe™i—lly in wiroung—D is more o˜vious th—n in rumerus @pigs ReD fY „—˜F PAF „his ˜one in m—les

other true se—lsF €rior to this studyD there h—ve never of gystophor— is —˜solutely longer —nd more ro˜ust

˜een —ny fossil rem—ins found —nd des™ri˜ed within gyE th—n in fem—lesD —nd the he—d is ™ompressed in — dorE

stophorin—eF „he new fossil post™r—ni—l ˜ones @SV indiE soEventr—l dire™tionD while in fem—les it is l—rger —nd

vidu—l ˜onesA ex—mined in this study show o˜vious diE more spheri™—l @™ontr—ry to €ho™in—eY „—˜F PAF „he r—tio

mension—l —nd morphologi™—l diversity —nd fell into two of the gre—test width of the he—d to the height in the

size ™l—sses th—t do not fit previously des™ri˜ed p—tterns m—le of gystophor— ™rist—t— is HFVSY in the fem—les it is

of individu—lD ontogeneti™D or sexu—l v—ri—tionsF €reE HFVHF hespite the t—ller˜one in — m—leD the length of the

viouslyD uoretsky @IWVU—D PHHIA studied sexu—l diE deltoid ™rest is equ—l to th—t in fem—lesD ˜ut the dist—l

morphism in the skullD m—ndi˜leD humerusD —nd femur p—rt of the deltoid ™rest in — fem—le ends —s — n—rrow †D

of ‚e™ent €ho™in—e —nd ™omp—red them to fossil m—teE while in — m—le it is widerD Elike @pigF QeAF „he he—d of

ri—lF „he finds of fossil gystophorin—e provided — unE the humerus of the m—le gF ™rist—t— @„—˜F PA is ˜ent

ique opportunity to study the sexu—l dimorphism in the ™—ud—lly to — gre—ter degree th—n th—t of fem—lesD indiE

lim˜ ˜ones —nd m—ndi˜les in extin™tD —s well —s livingD ™—ting pro˜—˜le sexu—l dimorphism —lsoF roweverD in

mem˜ers of the ƒu˜f—milyF „he following sexu—l differE the humerus of modern €ho™in—e this differen™e is not

en™es were identified @simil—r differen™es ™—n ˜e o˜E pronoun™edF

served in modern —nd fossil €ho™in—eAF „he foss— lo™—ted medioEdist—l to the he—d seen in

w—ndi˜le @pigs PfD gAF „he sexu—l dimorphism in ™—ud—l —spe™t @˜etween the lessertu˜er™le—nd the he—dA

gystophor— ™rist—t— is dis™erni˜le not just in size ˜ut in is deeperin m—les @—s in €ho™in—eAF „he enormousmeE

the different proportions of the r—mus —nd the ˜ody of di—l he—d of the tri™eps mus™le —rises from this foss— on

the m—ndi˜leF sn —dditionD the m—le retrom—ndi˜ul—r the medi—l side of the ne™k of the humerusD —nd inserts

sp—™e is even more elong—ted th—n in fem—lesD —s well onto the dors—l p—rt of the ole™r—non of the uln— @roE

—s in ˜oth sexes in other represent—tives of €ho™id—eD well IWQHY inglish IWUUAF „he —™tion of the tri™eps meE

with the ™oronoid ™rest less in™linedF „he ™oronoid proE di—lis mus™le is to extend the el˜ow joint @willeret —lF

™ess in m—les is mu™h lowerD is turned ™—ud—llyD —nd ends IWTRAF

—t —lmost the s—me level —s the ™ondyloid pro™ess @in sn fem—lesD —lthough the deltoid ™rest is ˜igger in

™ontr—st to other su˜f—miliesAF „he —ngul—r pro™ess is ™omp—rison to the rest of the ˜one @™ontr—ry to €ho™iE

more developed in fem—les th—n in m—lesY in ™ontr—stD n—eAD the deltoid tu˜erosityD whi™h is p—rt of the deltoid

the m—ndi˜ul—rnot™h in m—les is widerth—n in fem—lesF ™restD is more developed @simil—r to €ho™in—eAF „he musE

„he m—sseteri™ foss— in fem—les is —lso deeper —nd ˜etter ™ulospir—l grooveD where the ˜r—™hi—lis mus™le origiE

outlined th—n in m—lesF „he ™ondyloid —ngle @the posiE n—tes @rowell IWQHY €ieÂr—rd IWUIY „—r—soff IWUPY roE

tion of the ™ondyloid pro™ess in rel—tion to the —xis of w—rd IWUSY inglish IWUUAD is deeper in m—lesF sn fossilsD

the —lveol—r rowA of PS spe™imens were ex—minedD with this ™h—r—™ter is mu™h less pronoun™edD evidently —s —

the —ver—ge me—surements ˜eing SQë in fem—les —nd RHë result of we—r on the —v—il—˜le m—teri—lF

widdle wio™ene se—ls from the xorthern €—r—tethys QQQ

ƒexu—l dimorphism in the stru™ture of the di—phyE

sis is —sso™i—ted with v—rious degrees of development of

the v—stus intermedius —nd medi—lis mus™les in ™r—ni—l

—spe™t @see €ieÂr—rd IWUIX UQAD —nd of the mF —ddu™tor

™r—ni—lis @a —ddu™tor —nti™us in rowell IWQHA in ™—ud—l

—spe™tF e det—iled des™ription of the —™tions of these

mus™les ™—n ˜e found in uoretsky @PHHIAF

sn m—lesD the he—d of the femuris l—rger—ndthe

di—physis is less ™ompressedD while in fem—les it is the

reverseF „he —ver—ge width of the di—physis is sm—ller in

fem—lesD while the dist—l epiphysis is more developed in

m—lesF yne unique —nd very primitive ™h—r—™ter of gF

™rist—t— is th—t the lessertro™h—nterinm—les is well

developedD while in fem—les it is notF

sn ™ontr—st to €ho™in—eD the ne™k of the femur of

gF ™rist—t— is shorter —nd wider in fem—lesF „herefore @—s

in €ho™in—eAD the ne™k forms ne—rly — right —ngle with

the long —xis of the ˜one in fem—lesD ˜ut in m—les the

—ngle is gre—ter th—n WHëF

„he gre—ter tro™h—nter is wider —nd longer in

m—les th—n in fem—lesF „he dist—l p—rt of the gre—ter

tro™h—nter in fem—les termin—tes more sh—rply or

—™utely @˜eing †Esh—pedAD while in m—les it is frequently

rounded @ Esh—pedAF

„he gluteus medius —nd gluteus minimus mus™les

insert onto the ™r—ni—l side of the gre—ter tro™h—nterD

while the mF piriformis is —tt—™hed to the ™—ud—l sideF

„he —tt—™hment sites of these mus™les —re signifi™—ntly

more developed in m—les @rowell IWQHY €ieÂr—rd IWUIY

pigF R E ƒexu—l dimorphism of ‚e™ent gystophor— ™rist—t—F ruE row—rd IWUSY uoretsky PHHPAD —nd ™onsequentlyD the

merusD ™r—ni—l view of eA fem—le @ewxr IVRTTHD vFA

tro™h—nter —lso is more developed in m—lesF

—nd fA m—le @ewxr IVRTSWD vFAF w—le femur@ ƒxw

rowell @IWPVD IWQHAD €ieÂr—rd @IWUIAD —nd row—rd

SSHRIID ‚FA in gA ™r—ni—l —nd hA ™—ud—l viewsF

@IWUSA des™ri˜ed the tro™h—nteri™ foss— —s — pl—™e of

—tt—™hment forthe o˜tur—tor internus —nd externus musE

™lesD whi™h sh—re — ™ommon tendon of insertion with

sn fem—lesD the medi—l epi™ondyle is more ™omE

the superior —nd inferior gemelli mus™les @uoretsky 8

pressed —nd n—rrow in its l—ter—l p—rt @rowell IWQHY

ƒ—nders PHHPAF „he o˜tur—tor externus —nd gemelli musE

inglish IWUUAF et the s—me timeD the l—ter—l epi™ondyle

™les —re more developed in fem—les th—n in m—lesD resulE

is shorter —nd n—rrower th—n in m—lesD —s in €ho™in—eF

ting in — deeper —nd more ™losed tro™h—nteri™ foss— @—s in

„he entepi™ondyl—r for—men is —lw—ys present in ˜oth

€ho™in—eAF

sexesF

sn fem—les of gF ™rist—t—D the pl—nt—rfoss— —˜ove

„he sh—pe of the humer—l ™oronoid foss— of gysE

the l—ter—l ™ondyle is wider —nd deeper —nd is ˜ordered

tophor— depends on sexD with m—les h—ving — sh—rp triE

˜y — very thin edge of ˜oneF „he pl—nt—r foss— itself is

—ngul—r form —nd fem—les h—ving — roundedEtri—ngul—r

the pl—™e of origin of the pl—nt—ris —nd l—ter—l he—d of the

or semiErounded form @pigF ReAY its depth is somewh—t

g—stro™nemius mus™lesF elthough the p—tell—rsurf—™eis

gre—ter in fem—lesF l—rger in m—lesD the sizes of the ™ondyles —re —lmost

pemur@pigs RgD hY „—˜F TAF „he —˜solute length equ—l in ˜oth sexesF pordet—iled fun™tion—l —n—lysis of

of this ˜one in m—les of gF ™rist—t— is less th—n in feE these mus™les see row—rd @IWUSA —nd uoretsky @PHHIAF

m—les @in ™ontr—st to €ho™in—eAY howeverD the m—le feE por determining the sex using the humerus —nd

mur is more ro˜ust —nd widerF woreoverD in fem—les of femurin the ƒu˜f—mily gystophorin—eDthe following

gF ™rist—t— the —nteroposterior di—meter of the di—physis ™h—r—™ters ™—n ˜e usedX IA rumerus ± over—ll sizeY length

is gre—ter th—n in m—lesD ˜ut the le—st width of the of deltoid ™rest —nd width of its middle p—rtY depth —nd

di—physis in fem—les is lo™—ted in the middle of the sh—pe of the ™oronoid foss—Y depth of the foss— lo™—ted

sh—ftD while in m—les it is shifted to the proxim—l porE ™—ud—l to the medi—l side of the ne™k of the humerusD

tion of the sh—ftF dist—l to the lessertu˜er™le@simil—rto €ho™in—eA —nd PA

QQR uoretsky sFeF 8 ‚—hm—t ƒFtF

of the lesser tro™h—nterY —lmost equ—lEsized ™ondylesF pemur ± over—ll sizeY —nteroposterior width @adorsoE

ventr—l thi™knessA of di—physisY length —nd thi™kness of ytherspe™ifi™ simil—ritiesto gystophor— in™ludeX ™r—E

ne™kY length —nd width of gre—ter tro™h—nterY —nteroposE nioE™—ud—l depression of the humer—l he—dY short delE

terior di—meter of dist—l epiphysisY presen™e of lesser toid ™restY enl—rgement of the deltoid ™rest in the midE

tro™h—nterF dleY —nd presen™e of the entepi™ondyl—r for—menF

„he most reli—˜le ™h—r—™ters for sex determin—tion ƒimil—rities to wiroung— —reX sh—llow —nd wide

in gystophorin—e —reX in m—lesD the —˜sen™e of the menE intertu˜er™ul—r groove of humerusY sh—llowD ov—l ™oroE

t—l tu˜er™leY the height of the first in™isor —lmost equ—E noid foss— extending to the s—me level —s medi—l epi™onE

ling the dimensions of the ™—nineY width of the dist—l dyleY noti™e—˜le femor—l intertro™h—nteri™ lineY very

epiphysis on the humerusY —nd presen™e of the lesser deep inter™ondyl—r grooveF

tro™h—nter of the femurF sn —dditionD €—™hypho™— differs from these two

‡hile the fe—tures noted —˜ove —re ™h—r—™teristi™ gener— of the ƒu˜f—mily gystophorin—e ˜yX mu™h sm—lE

for —dult —nim—lsD they —re not ne—rly —s pronoun™ed in lersizeY position —nd sh—pe of the lessertu˜er™leof the

young —nd su˜—dult individu—lsF yver—llD the est—˜lished humerusY presen™e of — distin™t lip ˜etween the femor—l

differen™es —nd v—ri—tions of ™h—r—™teristi™s —re f—irly he—d —nd the ne™kY —nd deeper tro™h—nteri™ foss—F ƒpeE

™onst—ntD —llowing rese—r™hers to sep—r—te individu—l ™ifi™—llyD €—™hypho™— differs from gystophor— ˜y the

elements of the extremities —™™ording to sexF reverse rel—tionship ˜etween l—ter—l —nd medi—l epi™onE

dyles —nd from wiroung— ˜yX the presen™e of the enteE

pi™ondyl—r for—men of the humerusY shorter deltoid

™restY protruding lesser tro™h—nter of the femurY higher €—™hypho™— genF nF

position of the gre—ter tro™h—nterY shorter femor—l ne™k „ype spe™iesX €—™hypho™— ukr—ini™—Y widdle ƒ—rm—ti—n of kr—ine

—nd l—rger he—d ™omp—red to the size of the ˜oneF

itymologyX €—™hysD qreekD thi™k @referring to ˜one hypertroE

phyAY pho™—D v—tinD from the qreek phokeD se—l @fFAF

hi—gnosisX vesser tu˜er™le of humerus sm—llD roundD —nd loE €—™hypho™— ukr—ini™— genF n spF nF

™—ted dist—l to he—d —nd gre—ter tu˜er™leY he—d ™ompressed ™r—nio™—uE

pigs SEIHD „—˜s IEV

d—llyY intertu˜er™ul—r groove wide —nd sh—llowY medi—l epi™ondyle fl—tE

tenedD spre—ding from lower p—rt of entepi™ondyl—r for—menY ole™r—non

itymologyX ukr—ini™—D from v—tinized form of kr—ineF

foss— deep —nd n—rrowF

rolotypeX ‚ight humerusD xwxr E€ TREUHID widdle ƒ—rm—E

qre—ter tro™h—nter of femur slightly higher th—n he—dY tro™h—nE

ti—nF

teri™ foss— deepY intertro™h—nteri™ ™rest fl—tD wideD —nd thi™kD re—™hing

„ype vo™—lityX uhomutovo †ill—geD honetsk regionD xovo—zov

lesser tro™h—nterY he—d l—rgeD se—ted on distin™t lip @sh—ped like mushE

distri™tD kr—ineF

roomA —nd wide ne™kY minimum width of di—physis shifted proxim—llyY

‚—ngeX widdle ƒ—rm—ti—n @fess—r—˜i—n ƒt—geD IIFPEIPFQ w—A

˜ody of femurswollenD thi™kD huskyD he—vyD —nd p—™hyosteos™leroti™Y

deposits of the xorthern fl—™k ƒe— ‚egionD kr—ineF

™ondyles —lmost equ—l in sizeF

hi—gnosisX w—xim—l enl—rgement of humer—l deltoid ™rest loE

ƒ™—pul—rspine ends smoothlyY infr—spinousfoss— deeper—nd

™—ted in middleY deltoid ™rest shorter th—n ‰ of ˜one lengthY l—ter—l

widerth—n supr—spinousF

epi™ondyle shorter —nd lower th—n medi—l @in ™ontr—st to €F ™h—pskiiAD

wedi—l surf—™e of uln— fl—ttenedD not ™on™—veY ole™r—non short

does not re—™h proxim—l p—rt of medi—l epi™ondyleY medi—l epi™ondyle

—nd thi™kY r—di—l not™h deep —nd longF

extending more proxim—l th—n ™oronoid foss—Y entepi™ondyl—r for—men

slium thi™kY ili—™ ™rest not —vertedD —nd ex™—v—ted on its ventr—l

lo™—ted —˜ove ™oronoid foss—Y ™oronoid foss— ov—lF

surf—™e —s in other won—™hin—e —nd gystophorin—eY iliope™tine—l emiE

€roxim—l p—rt of femor—l gre—ter tro™h—nter wider th—n dist—lY

nen™e fl—ttenedY foss— for mF gluteus medius sh—llowY —uri™ul—r foss—

tro™h—nteri™ foss— n—rrowY lesser tro™h—nter well developedY he—d m—sE

deepY —l—r spine ro˜ustY ili—™ tu˜erosity —nd ™—ud—l dors—l ili—™ spine

siveD se—ted on distin™t lip on wide ne™kY supr—™ondyl—r foss— deepD

not very well developedY ™—ud—l dors—l is™hi—l spine rounded —nd wideD

wideD re™t—ngul—rF

not protrudingY —l—r spine ro˜ustY —™et—˜ulum sh—llowF

‚eferred ƒpe™imensX sn —ddition to the holotypeD the following

€oplite—l not™h of ti˜i— sh—llow —nd wideY ti˜i—l ™rest rounded

spe™imens were found in kr—ineX humeriX xwxr E€ TREPRT @vFAD in dorsomedi—l dire™tionY ti˜i—l tu˜erosity fl—ttenedY mus™ul—r groove

zunl—rY TREUHQ @vFAD TREUIQ @vFAD olot—y— f—lk—Y s™—pul—eX wideY grooves on dist—l end sh—llow —nd fl—ttenedF

xwxr E€D TRERUU @vFAD TREUHP @‚FAD olot—y— f—lk—Y r—diiX

ell ˜ones —re thi™kD huskyD —nd p—™hyosteos™leroti™F

xwxr E€ TRERVI @vFD imm—tureAD TRERVP @vFD imm—tureAD olot—y—

sn™luded spe™iesX sn the widdle ƒ—rm—ti—n @IPFQEIIFP w—A of the

f—lk—Y uln—eX xwxr E€ TREUHS @vFAD olot—y— f—lk—Y TREQVQ @‚FAD TRE xorthern fl—™k ƒe— littor—l regionD southern kr—ineD two new ™ystoE

UIH @‚FD possi˜ly from the s—me individu—l —s holotypeAD uhomutovoY phorine spe™ies —re re™ordedX €—™hypho™— ukr—ini™— —nd €F ™h—pskiiF

TREUIID @‚FD from the s—me individu—l —s ri˜ TREUIID not des™ri˜edAD TRE

‚—ngeX widdle wio™eneD widdle ƒ—rm—ti—n @fess—r—˜i—n ƒt—geY

UIP @‚FAD heltok—menk—Y innomin—t—X xwxr E€ TRERUW @vFAD TRE

IIFPEIPFQ w—AD the xorthern fl—™k ƒe— regionD kr—ineD i—stern €—r—E

QRV @vFAD olot—y— f—lk—Y femor—X xwxr E€ TREQSR@‚FAD olot—y— tethysF

f—lk—Y xwxr E€ TREITT @‚FAD qnylozu˜ovY TRERUI @vFAD olot—y—

f—lk—Y ti˜i—e —nd fi˜ul—eX xwxr E€ TRERUP @‚FD proxim—l fr—gmentAD

gomp—risonsF €—™hypho™— h—s ™h—r—™ters simil—r TRERUQ @vFAD TRERUV @vFD proxim—l fr—gmentAD olot—y— f—lk—Y verte˜r—X

xwxr E€ TREUHR @thor—™i™AD olot—y— f—lk—F to other represent—tives of the ƒu˜f—mily gystophorin—e

su™h —sX —˜sen™e of the mus™ulospir—l groove of the huE

hes™riptionF ƒ™—pul— @pigs SeEgY „—˜F IAX „he merusY the protruding femor—l intertro™h—nteri™ ™rest

extending more proxim—l th—n the he—dY the presen™e s™—pul—rspine ends smoothly —nd does not re—™hthe

widdle wio™ene se—ls from the xorthern €—r—tethys QQS

pigF S E ƒ™—pul— of €—™hypho™— ukr—iE

ni™— @xwxr E€D TRERUUD

vFA in eD dors—l fA ventr—l

—nd gD glenoid viewsF ƒ™—pul—

of €F ™h—pskii @xwxr E€

TREUHUA in hA dors—l —nd iA

ventr—l viewsF

verte˜r—l ˜orderD whi™h is p—rti—lly ˜roken in —v—il—˜le th—n the ™oronoid foss—F „he entepi™ondyl—r for—men

spe™imensF „he —™romion is not developed —nd does not is lo™—ted —˜ove the ™oronoid foss—D with — l—ter—llyE

re—™h the ventr—l —ngleF sn the ™ervi™—l regionD the infr—E lo™—ted wide w—llF „he ™oronoid foss— is sh—llowD ov—lD

—rti™ul—r tu˜erosity shows —s — long ridge ™onne™ted extends further proxim—lly th—n the l—ter—l epi™ondyleD

with — thi™k —nd distin™t mus™ul—rline on the infr—spiE —nd ends —t the s—me level —s the medi—l epi™ondyleF „he

nous foss—F „he ™or—™oid pro™ess —nd s™—pul—r tu˜erosE ole™r—non foss— is deep —nd n—rrowF „he ™omplete ˜one

ity —re not developed eitherF „he infr—spinous foss— is is extremely husky with — very p—™hyos™leroti™ ™ondiE

deeper—nd widerth—n the supr—spinousF„he dist—l ˜orE tionF

der of the supr—spinous foss— ends —s — str—ight lineF sn ‚—dius @pigs UeD fY „—˜F QAX „he r—di—l tu˜erosity

the infr—spinous foss— the ™—ud—l —ngle forms — thinD is very l—rgeD wideD fl—tD —nd does not protrude l—ter—llyF

round edge —t the verte˜r—l ˜orderF „he glenoid foss— „he groove for —ll tendons in™luding the tendon of the

is deep with — thin mushroomElike lipF „he ™omplete mF —˜du™tor polli™is longus is wide —nd sh—llowD where—s

˜one is very thi™k —t the l—ter—l ˜order —nd ˜e™omes the ridge for the mF extensor digitorum ™ommunis is

mu™h thinner —t the verte˜r—l ˜orderF only sh—llowF foth —v—il—˜le ˜ones ˜elong to su˜—dult

rumerus @pigs TeD fY „—˜F PAX „he intertu˜er™ul—r —nim—lsF

groove is sh—llow —nd wideF „he widest portion of the ln— @pigs UgEiY „—˜F RAX „he medi—l surf—™e of

deltoid ™rest is lo™—ted in the middleY the ™rest extends the ˜one is fl—ttened —nd not ™on™—veY the ole™r—non is

—˜out h—lf the length of the ˜oneD —nd smoothly desE short —nd thi™kD gr—du—lly ™onne™ting to the proxim—l

™ends to the ™ondyles —s — fl—tD —lmost invisi˜le ridgeF h—lf of the ˜oneF yn the l—ter—l surf—™eD there is ˜—rely

„he deltoid tu˜erosity is sm—ll —nd lo™—ted —t the middle —ny visi˜le prominen™e of the rugosity for ˜r—™hi—lis

of the deltoid ™rest proxim—l to the middle of the di—E mus™le insertionF ‚—ther th—n — foss— for origin of the

physisF „he lesser tu˜er™le is we—kly developedD roundD mF —˜du™tor polli™is longusD — protu˜er—n™e is presentF

—nd lo™—ted ™onsider—˜ly inferior to the gre—ter tu˜er™leD g—ud—l to the —rti™ul—r surf—™e is —nother deepD wideD

just dist—l to the he—dD whi™h is ™ompressed ™r—nio™—udE —nd short grooveY this groove is surrounded ˜y — ™ir™uE

—llyF „he mus™ulospir—l groove is —˜sentF „he l—ter—l l—r @sh—rpA ™restF yn the ˜one9s r—di—l —spe™tD the ™oroE

epi™ondyle is shorter —nd lower th—n the medi—lD —nd noid pro™ess only slightly protrudes forw—rd over the

does not re—™h the dist—l p—rt of the deltoid ™restF „he r—di—l not™hY the r—di—l not™h is deep —nd longF „he

medi—l epi™ondyle is fl—ttenedD spre—ding from the lower interosseous ™rest is swollenD forming — prominen™e th—t

p—rt of the entepi™ondyl—r for—menD extending higher protrudes only slightlyF „he he—d is not preservedF

QQT uoretsky sFeF 8 ‚—hm—t ƒFtF

pigF T E €—™hypho™— ukr—ini™— huE

merus @xwxr E€ TRE

PRTD vFA in eA l—ter—l —nd

fA medi—l viewsF €F ukr—iniE

™— femur@xwxr E€ TRE

ITTD ‚FA in iA ™—ud—l —nd

pA ™r—ni—l viewsF €F ™h—pskii

humerus @xwxr E€ TRE

SPQD vFA in gA l—ter—l —nd

hA medi—l viewsF €F ™h—pskii

femur@xwxr E€ TREUHTD

vFD reversedA in qA ™—ud—l

—nd rA ™r—ni—l viewsF

Characters OR „—˜F Q E we—surements @mmA of r—dii

€—™hypho™— ukr—in—F Absolute length 80.0; 85.0 of Width of diaphysis 12.5; 14.0 Width of distal epiphysis 23.0; 24.0 Width of articulation surface of distal epiphysis 16.0; 16.0 Width of articulation surface, disposition of medial from articulation cavity 9.0; 12.0 Lesser diameter of caput 13.0; 13.5 Least width of proximal epiphysis 10.5; 11.0 Greater diameter of caput 16.0; 16.5

Anteroposterior diameter of distal epiphysis 16.0; 16.0

Pachyphoca „—˜F R E we—surements @mmA of ulE Characters ukrainica chapskii n—eF nM OR Absolute length 2 113.0; 120.0 170.0; 155.0 Width of incisura trochlearis in upper part 4 12.3 10.0 - 13.5 14.0; 16.5 Width of incisura trochlearis in lower part 4 9.0 6.0 - 11.0 11.5; 12.0 Maximal width of middle part of diaphysis 4 14.7 13.0 - 18.5 18.5; 19.0 Width of bone at the level of incisura trochlearis 4 23.0 12.0 - 31.5 34.0; 35.0 Width of olecranon 4 35.3 30.5 - 40.0 45.5; 46.5 Transverse width of olecranon 4 13.3 12.0 - 14.0 16.5; 19.0

Maximal width of distal epiphysis 2 15.0; 17.5 19.5; 21.0

snnomin—te @pigs VgD hY „—˜F SAX es in other foss— fororiginof the mF pso—s m—jorF „he —l—rspine

won—™hin—e —nd gystophorin—eD the ilium is thi™k —nd does not protrude very f—rF „he iliope™tine—l eminen™e is

the ili—™ ™rest is not —verted —nd not ex™—v—ted on its —˜sentD —s is the pe™tine—l tu˜er™leF „he edges of the

ventr—l surf—™eF „he iliope™tine—l eminen™e is sm—llD siE —™et—˜ul—r foss— —re not r—ised —˜ove the pl—ne surf—™e

tu—ted higher th—n the proxim—l ˜order of the —™et—˜ul—r of the ˜oneY the —™et—˜ulum is ™ir™ul—r with — slightly

foss—F „he gre—ter is™hi—l not™h is slightly ™on™—veD —lE m—rked ™otyloid not™hF „he is™hium is thi™kD wide —nd

most str—ightF e sh—llow depression @foss—A for the mF ro˜ustY the is™hi—l spine is l—rgeD rounded —nd well deE

gluteus medius is lo™—ted on the medi—l —spe™t of the ili—l veloped for—tt—™hment of the mF ˜i™eps femorisF „he

wingY on its l—ter—l —spe™t is — deep —nd n—rrow —uri™ul—r pu˜is is not preservedF

widdle wio™ene se—ls from the xorthern €—r—tethys QQU

pigF V E snnomin—te ˜one in eA l—ter—l @aventr—lA —nd fA medi—l pigF U E €—™hypho™— ukr—ini™— r—dius @xwxr E€ TRERVID vFA in

@adors—lA views of ‚e™ent gystophor— ™rist—t— @ ƒxw eA dors—l —nd fA ventr—l viewsF €F ukr—ini™— uln—

SSHRIID `D ‚FAF snnomin—te ˜one of widdle wio™ene €—E @xwxr E€ TREUIHD ‚FA in gA medi—lY hA l—ter—lY —nd

™hypho™— ukr—ini™— @xwxr E€ TRERUWD vFA in gA l—ter—l iA ™r—ni—l viewsF €F ™h—pskii uln— @xwxr E€ TRESIVD

—nd hA medi—l viewsF snnomin—te ˜one of €F ™h—pskii ‚FA in pA medi—lY qA l—ter—lY —nd rA ™r—ni—l viewsF

@xwxr E€ TRESPSD ‚FA in iA l—ter—l —nd pA medi—l

viewsF e˜˜revi—tionsX —D —™et—˜ulumY —fD —uri™ul—r foss—Y

—nD —™et—˜ul—r not™hY ™disD ™—ud—l dors—l ili—™ spineY ™rdisD pemur @pigF TiD pY „—˜F TAX „he gre—ter tro™h—nter

™r—ni—l dors—l ili—™ spineY ™rvisD ™r—ni—l ventr—l ili—™ spineY extends proxim—lly slightly higher th—n the he—dY its

gisnD gre—ter is™hi—l s™i—ti™ not™hY ilD iliumY iltu˜D ili—™ tuE

proxim—l p—rt is somewh—t wider th—n the dist—lF „he

˜erosityY ipeD iliope™tine—l eminen™eY isD is™hiumY isspD isE

tro™h—nteri™ foss— is deepD n—rrow —nd open proxim—llyD ™hi—l spineY istD is™hi—l tu˜erosityY mgmD mF gluteus medE

re—™hing the dist—l h—lf of the gre—ter tro™h—nterF „he iusY ofD o˜tur—tor for—menY ogD o˜tur—tor grooveY pD pu˜isY

rfD mF re™tus femoris —tt—™hmentY spD symphysis pu˜isF fl—tD wide —nd thi™k intertro™h—nteri™ ™rest re—™hes the

lesser tro™h—nterD whi™h serves for insertion of the mF

he—d @orslightly higherth—n the he—dAF „he femor—l iliopso—sF „he lesser tro™h—nter is very visi˜leD thi™kD —nd

he—d is l—rgeD rel—tive to the ˜one9s m—ssD —nd se—ted not sep—r—ted from the ™restY the lesser tro™h—nter

on — very shortD thi™kD —nd wide ne™kY ˜etween the he—d re—™hes ˜elow the dist—l ˜order of the gre—ter tro™h—nter

—nd ends —t the s—me level —s the dist—l ˜order of the —nd ne™k is — distin™t lipF „he sm—llest width of the

QQV uoretsky sFeF 8 ‚—hm—t ƒFtF

Characters P. ukrainica P. chapskii thigh upon the pelvisD —nd —ddu™ts the femurY uoretsky OR 8 ƒ—nders PHHPAF

Length from center of acetabulum to iliac crest 52.5 70.5

B

os ™ox— is —lmost

Length of pubic from center of acetabulum 61.0 - „he gre—ter is™hi—l not™h of the Width of level of iliac crest 45.0 53.0 str—ightF

Width of pubic 60.0; 67.0 44.0 B

Thickness of pubic 11.0; 13.5 14.0 e deep —nd n—rrow —uri™ul—r foss— is present for

mF pso—s m—jor @— flexorof the hip jointAF

Ratio insertion of the

1 : 3 117.6 133.0 B „he is™hi—l spine is l—rgeD roundedD —nd well

developed for—tt—™hment of the mF ˜i™eps femoris @—n „—˜F S E we—surements @mmA of innomin—te of €—™hypho™—F

extensorof the hip jointAF

„hereforeD it ™—n ˜e ™on™luded th—t this —nim—l

di—physis is shifted tow—rd the proxim—l end of the

w—s well —d—pted for terrestri—l lo™omotion due to the

˜oneF „he supr—™ondyl—r foss— is lo™—ted —˜ove the presen™e of these mus™ul—r —tt—™hmentsF

medi—l ™ondyle —nd is deepD wideD —nd re™t—ngul—rY the

™ondyles —re —lmost equ—l in sizeF

€—™hypho™— ™h—pskii genF nF spF nF

„i˜i— @pigsF WeEgY „—˜F UAX „he two ™ondyles —re

pigs SEIHD „—˜s ID PD REW we—kly ™on™—ve in their ™entersD shortened —nd ov—lF

„he inter™ondyloid eminen™es —re we—k —nd only

itymologyX x—med in honorof the l—te uF uF gh—pskii @of the

slightly r—ised —˜ove the two l—ter—lD wellEdeveloped oologi™—l snstituteD ƒtF €eters˜urgD ‚ussi—A in re™ognition of his nuE

merous ™ontri˜utions to the study of fossil —nd modern true se—lsF ˜orders of the ™ondylesF „he poplite—l not™h is sh—llow

rolotypeX veft femurD xwxr E€ TREUHTD widdle ƒ—rm—ti—nF

—nd wideD ˜ut well m—rkedF „he ti˜i—l ™rest is rounded

„ype vo™—lityX heltok—menk—D vill—ge on heltenk—y— ‚iverD

in — dorsomedi—l dire™tionF yn the ventr—l side of the hnepropetrovsk regionD epostolovo distri™tD kr—ineF

ti˜i—D the ti˜i—l tu˜erosity is fl—ttened —nd not well ‚—ngeX widdle ƒ—rm—ti—n @fess—r—˜i—n ƒt—geD IIFPEIPFQ w—A

deposit of the xorthern fl—™k ƒe— ‚egionD kr—ineF m—rkedF „he mus™ul—r groove is fl—t —nd wideY the

hi—gnosisX wedi—l epi™ondyle does not re—™h ™oronoid foss— @in

grooves on the dist—l end —re sh—llow —nd fl—ttenedF

™ontr—st to €F ukr—ini™—AY ™oronoid foss— wideD not well outlinedD does

his™ussionF „he innomin—te ˜one of €—™hypho™— not re—™h proxim—l end of l—ter—l epi™ondyleF

ukr—ini™— h—s sever—l very primitive ™h—r—™teristi™sD su™h qre—ter tro™h—nter of femur is o˜liqueY its dist—l p—rt is wider

th—n the proxim—lY tro™h—nteri™ foss— is wide —nd re—™hes middle of —sX

gre—ter tro™h—nterY lesser tro™h—nter is wideD not sep—r—ted from interE

B

„he femor—l lesser tro™h—nter is presentD for inE

tro™h—nteri™ ™restY ™ondyles fl—tD sm—llD widely sp—™edY inste—d of pl—nt—r

sertion of the mF iliopso—s @whi™h flexes —nd l—ter—lly

foss—D pl—nt—r eminen™e presentY supr—™ondyl—r foss— h—s thi™k protuE

rot—tes the femurD tilts the pelvis forw—rdD flexes the ˜er—n™eF

Pachyphoca Cystophora cristata Characters ukrainica chapskii female male n M OR n M OR n M OR Absolute length 3 80.3 78.0 - 82.5 120 6 131.0 130.0-132.0 3 132.4 131.0-133.5 Medial length 2 74.5; 78.0 - 6 119.5 119.0-120.0 3 121.2 119.0-122.6 Lateral length 3 75.3 72.5; 78.0 11 6 115.7 115.0-116.0 3 117.6 115.0-120.0 Length of medial condyle 3 15.3 14.0 - 18.0 22.5 6 22.2 22.0-26.0 3 24.3 22.1- 28.0 Length of lateral condyle 3 16.0 15.0 - 17.0 24 6 28.4 27.6-29.0 3 28.9 27.5- 30.0 Length of greater trochanter 2 22.0; 26.0 37 6 39.3 39.0-40.0 3 42.0 41.0- 43.5 Intertrochanter length 3 33.5 33.0 - 34.0 39 6 54.8 54.0-55.0 3 55.8 54.5- 58.0 Height of head 2 16.6; 16.5 - 6 25.7 25.0-26.5 3 27.2 26.1- 28.0 Height of articular area of patella surface 3 16.0 14.5 - 18.0 20.5 6 31.2 30.0-32.0 3 31.2 30.0- 33.0 Width of proximal epiphysis 2 36.5; 39.0 - 6 35.8 65.0-66.0 3 67.8 65.8- 69.0 Width of distal epiphysis 3 38.5 36.5 - 41.0 56 6 74.5 73.5-75.0 3 72.5 69.6- 75.0 Width of condyles 3 33.9 33.0 - 37.0 58 6 60.9 57.0-63.0 3 57.9 56.1- 61.0 Width of greater trochanter 2 15.5; 17.0 24 6 25.0 23.0-27.0 3 26.0 23.6- 30.0 Width of head 2 16.5; 16.5 - 6 26.2 24.0-30.5 3 28.4 25.5- 34.2 Width of diaphysis 3 24.3 22.0 - 26.0 33.5 6 33.1 32.0-34.0 3 34.4 33.5- 35.0 Anteroposterior thickness of diaphysis 3 14.3 12.0 - 17.0 21.5 6 25.0 20.5-27.0 3 24.6 19.6- 29.2 Thickness of medial condyle 3 17.5 17.0 - 18.0 27 6 26.6 24.0-27.5 3 30.9 25.0- 36.0 Thickness of lateral condyle 2 20.0; 22.5 30 6 37.0 35.0-39.0 3 37.8 35.8- 40.0 Distance between condyles 3 9.5 8.5 - 11.0 15.5 6 18.2 17.0-20.0 3 17.5 16.0- 19.0 Diameter of neck 2 16.0; 17.0 - 6 20.8 15.0-23.0 3 27.0 26.4- 28.0 Sex ratio 09:11 3 41.6 37.8 - 47.4 36.7 6 41.9 39.8-43.2 3 43.0 41.0- 44.0 20:01 2 20.0; 20.6 - 6 15.9 14.2-17.3 3 20.5 19.8- 21.4

13:06 2 77.3 65.9 6 63.6 56.8-67.7 3 62.0 57.1- 69.7 „—˜F T E we—surements @mmA of femur—F

widdle wio™ene se—ls from the xorthern €—r—tethys QQW

pigF W E €—™hypho™— ukr—ini™—

@xwxr E€ TRERUQD vFA

ti˜i— in eA ™—ud—lY fA ™r—E

ni—l —nd gA proxim—l

viewsF hA €F ™h—pskii ti˜i—

@xwxrE€ TRESPHD vFD

imm—tureA in ™—ud—l viewF

€F ™h—pskii fi˜ul—

@xwxr E€ TRESPID vFD

imm—tureD from the s—me

individu—l —s TRESPHA in

iA ™—ud—l —nd pA ™r—ni—l

viewsF

Pachyphoca xwxr E€ TRESPI @vFD imm—tureD from the s—me individu—l —s TRE

verte˜r—X xwxr E€ TRESPV @lum˜—rAD heltoE

Characters ukrainica chapskii SPHAD heltok—menk—FY

s—™rumX xwxr E€ TRESPUD heltok—menk—Y swt™ Pnd

nM OR k—menk—Y

X xwxr E€ TRESIUD heltok—menk—F Width of proximal epiphysis 3 33.2 32.0 - 35.0 40.0 ph—l—nge Height of proximal epiphysis 3 20.0 18.0 - 22.0 27.0

Width of distal epiphysis 2 20.5; 21.5 31.5

hes™riptionF ƒ™—pul— @pigs ShD iY „—˜F IAX „he

„—˜F U E we—surements @mmA of ti˜i—eF s™—pul—r spine ends smoothlyD re—™hing the verte˜r—l

˜order —s — sm—llD —lmost fl—t ridgeY the verte˜r—l ˜order

is ˜roken on the —v—il—˜le spe™imenF „he —™romion is

‚eferred ƒpe™imensX sn —ddition to the holotypeD the following

present —s — sm—ll hookD ending —t the s—me level —nd

spe™imens were found in kr—ineX humeriX xwxr E€ TREUHV @‚FD

height —s the s™—pul—rspineF

dist—l endAD TRESPQ @vFAD heltok—menk—Y s™—pul—X xwxr E€D TREUHU

sn the ™ervi™—l regionD the infr—E—rti™ul—r tu˜erosE

@vFAD heltok—menk—Y uln—eX xwxr E€ TRESIV @‚FAD TRESIW @‚FAD

ity is present —s — thi™kD wide protu˜er—n™eF „he musE heltok—menk—Y innomin—teX xwxr E€ TRESPS @‚FAD heltok—menE

k—Y ti˜i—X xwxrE€ TRESPH @vFD imm—tureAD heltok—menk—Y fi˜ul—X ™ul—r line on the infr—spinous foss— is pronoun™ed —s —

QRH uoretsky sFeF 8 ‚—hm—t ƒFtF

longD very well developed ridge @˜igger —nd stronger ™hin—e —nd gystophorin—e @simil—r to €F ukr—ini™—AF „he

th—n in €F ukr—ini™—AF „he infr—spinous foss— is deep iliope™tine—l eminen™e is wide —nd not pronoun™edY the

pe™tine—l line is h—rdly visi˜leF yn the medi—l —spe™t of —nd wider th—n the supr—spinous foss—F sn ™ontr—st to

the ilium is — sh—llow depression @™omp—red to size of €F ukr—ini™—D the ™or—™oid pro™ess is thi™kD shortD —nd

the ˜oneA forthe insertionof the mF gluteus medius @this wideF

is mu™h sm—ller—nd sh—llowerth—n the s—me foss— of €F „he dist—l ˜order of the supr—spinous foss— ends

ukr—ini™—AF yn the l—ter—l —spe™t of the ili—l wing is —s — str—ight line @—s in €F ukr—ini™—AY the ™—ud—l —ngle is

lo™—ted — deep —nd wide foss— forinsertionof the mF not preservedF „he glenoid foss— is sh—llow @deep in €F

pso—s m—jorF „he —l—r spine is roundedD ro˜ustD —nd ukr—ini™—AD without — pronoun™ed lipD —nd without —

˜order @in ™ontr—st to €F ukr—ini™—AF turned posteriorlyF „he edges of the —™et—˜ulum —re

„he ™omplete ˜one is very thi™kD he—vyD —nd mu™h r—ised high —˜ove the pl—ne surf—™e of the ˜one @in ™onE

˜igger—nd thi™kerth—n in €F ukr—ini™—F tr—st to €F ukr—ini™—AY the —™et—˜ulum is ™oni™—l @— priE

rumerus @pigs TgD hY „—˜F PAX „he l—ter—l epi™onE mitive ™h—r—™terAD with — deep —nd wellEm—rked ™otyloid

dyle is mu™h higher—nd widerth—n the medi—lF „he not™hF e˜ove the —™et—˜ulumD there is — deep —nd n—rE

middle p—rt of the tro™hle— is r—ised w—veElike over row depression for the insertion of the mF re™tus femoris

the ™oronoid foss—F „he sm—llD ov—l entepi™ondyl—r for—E @in ™ontr—st to €F ukr—ini™—D where there is — ˜igD fl—tD —nd

men h—s — very wide ˜ridge over the medi—l epi™ondyleD wide depressionAF „he pu˜is is roundedF

—nd extends lowerth—n ˜oth the ™oronoidfoss— —nd the pemur @pigs TqD rY „—˜F TAX „he gre—ter tro™h—nE

l—ter—l epi™ondyleF ter is very o˜liquely oriented —long the ˜one9s —xisY its

„he medi—l epi™ondyle is fl—ttenedD spre—ding dist—l p—rt is wider th—n the proxim—l —nd ends in —

from the lower p—rt of the entepi™ondyl—r for—menD n—rrow †Esh—peF „he tro™h—nteri™ foss— is deep —nd

—nd it does not re—™h the ™oronoid foss—F „he medi—l wideD re—™hes the middle of the gre—ter tro™h—nter9s

epi™ondyle is lower th—n the ™oronoid foss—Y the ™oroE lengthD —nd h—s — sh—llow proxim—l ˜orderF „he fl—tD

noid foss— is sh—llowD wide —nd not well defined @aoutE wideD —nd thi™k intertro™h—nteri™ ™rest re—™hes the

linedAY it extends f—r proxim—llyD ˜ut it does not re—™h we—kly developed lessertro™h—nterDwhi™h is wide —nd

the proxim—l end of the l—ter—l epi™ondyleF not sep—r—ted from the intertro™h—nteri™ ™restF „he ™onE

„he ole™r—non foss— is deepD longD —nd n—rrowD dyles —re fl—t —nd sm—llD ™omp—red to the size of the

se—ted tr—nsversely to the ˜one —xisF ƒitu—ted on the ˜oneD —nd very widely sp—™edF gonsider—˜le swelling

—nteriorEmedi—l side of the humerus is — thi™k protruE of the ˜one is seen —t the lo™—tion of the pl—nt—rfoss—F

sionD lo™—ted medi—l to the tro™hle— —nd —dj—™ent to the supr—™ondyl—r foss— is not pronoun™edD ˜ut does „he

ole™r—non foss—F „he ˜one is ˜igD strongD —nd he—vy like h—ve — thi™k protu˜er—n™eF „he sm—llest width of the

— stoneF di—physis is shifted tow—rd the proxim—l end of the

ln— @pigs UpErY „—˜F RAX „he medi—l surf—™e is ˜oneF „he ™ondyles —re —lmost equ—l in sizeF

fl—ttened —nd not ™on™—ve @—s in €F ukr—ini™—AF „he oleE „i˜i— —nd fi˜ul— @pigs WhEpY „—˜F UAX „he two

™r—non is shortD thi™kD —nd forms — hook —t the ™—ud—l ti˜i—l ™ondyles —re shortened —nd ov—lY the ti˜i—l ™rest

endF „he rugosity for the ˜r—™hi—lis mus™le insertion h—s is well m—rked —nd developedF „he inter™ondyloid emiE

— very well developed prominen™eF yn the l—ter—l surE nen™e is not developedD only slightly r—ised —˜ove the

f—™eD the foss— forthe mF —˜du™tor polli™is longus is preE two l—ter—l ™ondylesD with h—rdly noti™e—˜le ˜ordersF

sentF enother n—rrow —nd long groove is lo™—ted ™—ud—l „he poplite—l not™h is sh—llow —nd wideD ˜ut not well

to the —rti™ul—r surf—™eY this groove is not surrounded ˜y m—rkedF „he ti˜i—l ™rest is well pronoun™ed —nd

— sh—rp ™restD ˜ut very smoothly tr—nsitions into the rounded in the dorsomedi—l dire™tionF yn the ventr—l

di—physisD re—™hing the uln—r protu˜er—n™e @in ™ontr—st side of the ti˜i—D the ti˜i—l tu˜erosity is fl—ttenedD wideD

to the uln— of €F ukr—ini™—AF „he tro™hle—r not™h is fl—tE tri—ngul—rD —nd extends —long the —xis of the ˜oneF „he

tened —nd not visi˜le on the r—di—l —spe™t of the ˜oneF mus™ul—r groove is deep —nd wideY grooves on the dist—l

„he ™oronoid pro™ess gre—tly protrudes forw—rdD —t —lE end —re sh—llow —nd fl—ttenedF „he fi˜ul— is without

most WHëD over the r—di—l not™hY the r—di—l not™h is deep ™ondylesD ˜ut h—s — wellEdeveloped interosseous ™restF

—nd longF „he —n™one—l pro™ess is very l—rgeD wideD —nd foth ˜ones pro˜—˜ly ˜elong to the s—me individu—l —nE

protrudingF „he uln—r tu˜erosityD situ—ted on the middle im—lF

of the di—physisD re—™hes the rugosity for the tendonF †erte˜r— @pigs IHhEpY „—˜F VAX „he ˜ody of the

„he ˜igD longD —nd wide interosseous ™rest is lo™—ted lum˜—r verte˜r— is fl—ttened in the dorsoEventr—l dire™E

on the middle of the di—physisF „he he—d —nd styloid tion —nd he—rtEsh—pedY the verte˜r—l for—men is ov—l in

pro™ess —re preservedF sh—peF „he spinous pro™ess is very short —nd wideD while

the tr—nsverse pro™esses —re dire™ted ™r—nioEventr—llyF snnomin—te @pigs ViD pY „—˜F SAX „he ilium is thi™k

„he ™—ud—l —rti™ul—r pro™ess is very short —nd roundedY —nd fl—ttenedD with the ili—™ ™rest slightly —verted —nd

the m—mmill—ry pro™ess is thi™kD wideD —nd shortF „he not ex™—v—ted on its ventr—l surf—™e —s in other won—E

widdle wio™ene se—ls from the xorthern €—r—tethys QRI

pigF IH E „hor—™i™ verte˜r— of €—™hyE

pho™— ukr—ini™— @xwxr

E€ TREUHRA in eA l—ter—l —nd

fA dors—l viewsF vum˜—r

verte˜r— of €F ™h—pskii

@xwxr E€ TRESPVA ™—uE

d—lY —nd iA l—ter—l viewsF ƒ—E

™rum of €F ™h—pskii

@xwxr E€ TRESPUA in pA

dors—lY qA ventr—lY —nd rA

™r—ni—l viewsF

gl—disti™ en—lysis ™r—ni—l —rti™ul—r surf—™e is roundedD like — short protuE

˜er—n™eF

„he d—t— m—trix for the RV in™luded ™h—r—™ters of

ƒ—™rum @pigs IHqEsY „—˜F WAX „he s—™rum ™onsists

€ho™id—e is shown in „—˜F IHF sn this m—trixD H desigE of R fused verte˜r—e —s in other ™ystophorines @with IPS

n—tes the most primitive st—te —mong the t—x— studiedY IE mm —˜solute length —nd UP mm wideAF „he s—™r—l proE

PD derived st—tesY cD unknown or missing d—t—F ƒome montory is —lmost fl—t —nd not pronoun™edF roweverD

™h—r—™ters h—ve the opposite pol—rity to th—t of fert— the —l— is thi™k —nd the —nterior surf—™e of the first ™enE

—nd ‡yss @IWWRA —nd furns —nd p—y @IWUHAD while some trum is lower th—n the wings of the s—™rum @—s in mirE

™h—r—™ters h—ve the s—me pol—rity —s th—t of gh—pskii oungini —nd mon—™hinesD ˜ut in ™ontr—st to ™ystophorE

@IWURAF por— det—iled dis™ussion of pol—ritiesof the ines —nd pho™inesD where the —l— is on the s—me level —s

™h—r—™tersD see uoretsky —nd qrigores™u @PHHPAF the ™entrumY entoniuk IWUWAF „he m—ximum width

—™ross the wings is SUFT 7 of the length of the s—™rumD ƒkull

whi™h —™™ording to entoniuk @IWUWA is — miroungine IF w—stoid pro™essX @HA prominen™e l—ter—l to —uditory ˜ull— not

strongly pronoun™edY @IA pronoun™edF ™h—r—™terD —s the per™ent—ge in ™ystophorines should

PF w—stoid pro™essX @IA n—rrow @width of the pro™ess less th—n

˜e less th—n RH7F yn ƒP —re lo™—ted thi™kD wideD —nd

length of pro™ess itselfAY @HA wide @gh—pskii IWURXQHIY in ™ontr—st to

wellEdefined tr—nsverse pro™essesF „he medi—n s—™r—l

fert— —nd ‡yss IWWRXRVAF

™restD whi™h is formed ˜y the spinous pro™essesD is QF w—xill—X @HA h—s very pronoun™ed ™onvexity —nterior to the

fused ˜etween ƒI —nd ƒPF ell spinous pro™esses ˜e™ome or˜itsY @IA short ™on™—vityY @PA long ™on™—vity @gh—pskii IWURY in ™onE

tr—st to fert— —nd ‡yss IWWRXRTAF progressively sm—ller from ƒI to ƒRD with immenseD

RF enterior p—l—t—l for—min—X @HA f—intly m—rked @furns —nd p—y

thi™k ˜—sesF hors—l s—™r—l for—min— —re sm—ller th—n

IWUHXUPAY @IA ov—l —nd sh—llowY @PA round —nd deepF

ventr—lF

SF snteror˜it—l sp—™eX @HA less th—n PSFH7 of width of skull —™ross

e™™ording to the sexu—lly dimorphi™ ™h—r—™ters of m—stoidsY @IA less th—n QHFH7D ˜ut equ—l to orgre—terth—n PSFH7 of

s—™r—l ˜ones des™ri˜ed ˜y q—djiev @IWVPAD this p—rti™ul—r width of skull —™ross m—stoidsY @PA equ—l to or gre—ter th—n QHFH7 of

width of skull —™ross m—stoids @furns —nd p—y IWUHXQUHY gh—pskii ˜one should ˜elong to —n —dult m—le ˜e™—use its first

IWURXPWWAF ventr—l for—men is rounded —nd the l—ter—l w—ll is very

TF tugul—r pro™essX @HA well developedY @IA poorly developedF

thi™kF „he ˜one is p—rti—lly rolledD ˜ut it is very distin™E

UF ‚ostrumX @HA elong—teY @IA shortD ™omp—red with skullF

tive —nd unusu—lF „he ˜ody of the s—™rum is shortD wideD VF hi—meter of infr—or˜it—l for—menX @HA less th—n di—meter of

—nd very p—™hyosteos™leroti™F —lveolus of m—xill—ry ™—nineY @IA —pproxim—tely equ—l to di—meter of

QRP uoretsky sFeF 8 ‚—hm—t ƒFtF

Pachyphoca PQF f—s—l ™ingulum of post™—nine teethX @HA well developedY @IA Characters not developedF

ukrainica chapskii

Height of centrum 21.0 27.5 PRF xum˜er of —ddition—l ™usps of premol—rsX @HA twoY @PA more Width of centrum 20.0 29.0 th—n twoF

Thickness of centrum 17.0 33.0 PSF €remol—rsX @HA se—ted p—r—llel to —xis of tooth rowY @IA se—ted

o˜liquelyF

PTF pper in™isorsX @HA —rr—nged in ™urved lineY @IA —rr—nged in

„—˜F V E we—surements @mmA of lum˜—r verte˜r—eF

str—ight lineF PUF pper in™isorsX @HA third l—rger th—n se™ond @€ho™in—eAY @IA

Characters Pachyphoca chapskii se™ond l—rger th—n first @won—™hin—eAY @PA in™isors equ—l in size to

Absolute length 125.0 ™—nine @gystophorin—eA @unordered ™h—r—™terAF B€ho™in—e h—ve IH inE ™isors @Q upperAY won—™hin—e h—ve V in™isors @P upperAY —nd gystophorE Hight of centrum 25.0 in—e h—ve T in™isors @I upperAF

Width of centrum 25.0

Thickness of centrum 25.0 rumerus

Thickness of 4-th vertebra 19.0 PVF vesser tu˜er™leX @HA pronoun™edY @IA not pronoun™ed @in

Length of ala 46.0 ™ontr—st to fert— —nd ‡yss IWWRX SPAF

Width of ala 23.5 PWF „ro™hle—r ™restX @HA r—ised in — w—veElike sh—pe over ™oroE

noid foss—Y @IA r—ised —r™hElike over ™oronoid foss—Y @PA not sep—r—ted

„—˜F W E we—surements @mmA of s—™rumF from ™oronoid foss— ˜y — distin™t lipF

QHF vesser tu˜er™le —nd he—dX @HA equ—l in height or tu˜er™le

insignifi™—ntly higher th—n he—dY @IA tu˜er™le very mu™h higher th—n

—lveolus of m—xill—ry ™—nineY @PA gre—ter th—n di—meter of —lveolus of

he—dY @PA tu˜er™le lower th—n he—d @unordered ™h—r—™terAF

m—xill—ry ™—nineF

QIF vesser tu˜er™leX @HA roundedY @IA extended —long the ˜one9s

WF enteroposterior length of —uditory ˜ull—eX @HA gre—ter th—n

—xisY @PA ov—lF

dist—n™e ˜etween themY @IA less th—n dist—n™e ˜etween themY @PA —˜out

QPF re—dX @HA medioEl—ter—lly ™ompressedY @IA roundY @PA fl—tE

equ—l to dist—n™e ˜etween them @unordered ™h—r—™terA @furns —nd p—y

tened proximoEdist—llyF

IWUHXQVPY g—hpskii IWURXQHHAF

QQF heltoid ™restX @HA m—ximum enl—rgement is in its proxim—l

p—rtY @IA neither p—rt noti™e—˜ly enl—rgedY @PA m—xim—l enl—rgement is in

w—ndi˜le

its middle p—rtF

IHF ƒymphyse—l p—rtX @HA ™ontinues —t le—st to the middle of the

QRF heltoid ™restX @HA shorter th—n oneEh—lf length of the ˜oneD

—lveolus of pQY @IA re—™hes only the —lveolus of pPY @PA re—™hes only the

™onfined to the proxim—l h—lf of the ˜oneY @IA longer th—n oneEh—lf

—lveolus of pIF

length of the ˜one ˜ut not re—™hing ™oronoid foss—Y @PA re—™hes ™oroE

IIF v—ter—l outline of symphyse—l regionX @HA squ—reD symphysis

noid foss— @in ™ontr—st to fert— —nd ‡yss IWWRX SPAF

thinY @IA roundedD symphysis thi™kY @PA str—ightD symphysis thi™kF

QSF goronoid foss—X @HA deepY @IA sh—llowF

IPF ghin prominen™eX @HA pronoun™edY @IA —˜sent or we—kly

QTF re—d —nd tro™hle—X @HA he—d wider th—n tro™hle—Y @IA he—d

outlinedF

—lmost equ—l in width to tro™hle—Y @PA tro™hle— wider th—n he—d @in

IQF ghin prominen™eX @HA extends from the —nterior or posterior ™ontr—st to fert— —nd ‡yss IWWRX SQAF

—lveolus of pP to the posterior or —nterior —lveolus of pRY @IA extends

from the —nterior —lveolus of pP to —nterior —lveolus of pQY @PA extends pemur

from the —nterior —lveolus of pP to posterior —lveolus of mI @unordered QUF vesser tro™h—nterX @HA presentY @IA —˜sent @fert— —nd ‡yss

™h—r—™terAF IWWRX SRAF

IRF w—ximum height of ˜ody of m—ndi˜leX @HA situ—ted ˜etween QVF gondylesX @HA different in sizeY @IA simil—r in sizeF

pP —nd pQY @IA situ—ted in the middle or —t the posterior portion of pPY QWF ipiphysesX @HA dist—l epiphysis widerth—n proxim—l˜y oneE

@PA situ—ted ˜etween —lveoli of pREmI orposteriorto—lveolus of mI fourth to oneEfifthY @IA widths of proxim—l —nd dist—l epiphyses —˜out

@unordered ™h—r—™terA @uorestky —nd ‚—y IWWRAF equ—lY @PA proxim—l epiphysis wider th—n dist—l oneF

ISF hi—stem—t— —nd tooth —lveoliX @HA —lveoli sm—llD with equ—l RHF ƒh—ftX @HA minimum width less th—n or—˜out equ—l to twoE

di—stem—t—Y @IA —lveoli round —nd l—rgeD with equ—l di—stem—t— ˜etween thirds width of proxim—l epiphysisY @IA minimum width more th—n twoE

themY @PA —lveoli sh—llowD —nd di—stem—t— unequ—l @unordered ™h—r—™E thirds width of proxim—l epiphysisF

terAF RIF sntertro™h—nteri™ ™restX @HA well developedY @IA —˜sent or

ITF elveoli of pR —nd mIX @HA —lveoli simil—rin sizeY @IA —lveolus poorly developedF

of pR sm—llerth—n —lveoli of mIY @PA —lveolus of pR l—rgerth—nthe RPF sntertro™h—nteri™ ™restX @HA re—™hes lower th—n he—dY @IA

—lveolus of mI @unordered ™h—r—™terAF shortD ends on s—me level —s dist—l edge of he—d or fove— ™—pitisF

IUF ‚etrom—ndi˜ul—r sp—™eX @HA elong—tedY @IA shortF RQF re—dX @HA roundY @IA fl—ttened in proximoEdist—l dire™tionY

@PA ™ompressed in medioEl—ter—l dire™tionF

„eeth RRF snter™ondyl—r —re—X @HA n—rrowD deepY @IA wideD sh—llowF

IVF xum˜erof in™isorsX@HA QGPY @IA PGPY @PA PGI @gh—pskii IWURX RSF qre—ter tro™h—nterX @HA m—ximum width in its middle p—rtY

PVWY in ™ontr—st to furns —nd p—y IWUHX QVHAF @IA m—ximum width in its proxim—l p—rt @uoretsky IWVUAF

IWF ‚oots of post™—nine teeth @€Dp P ± €Dp RAX @HA one rootD RTF re—d —nd gre—ter tro™h—nterX @HA ˜oth re—™h s—me levelY @IA

divided p—rti—lly —t the ˜—seY @IA two @fert— —nd ‡yss IWWRX SIAF gre—ter tro™h—nter higher th—n he—dY @PA he—d higher th—n gre—ter troE

PHF growns of post™—nine teethX @HA singleE™uspedY @IA multiE ™h—nterF

™usped @revers—l to primitive ™onditionA @fert— —nd ‡yss IWWRX SIAF RUF xe™kX @HA longD slenderY @IA shortD wideF

PIF ‚el—tive dimensions of post™—nine teeth to the size —nd RVF ƒh—ftX @HA minimum width in its proxim—l p—rtY @IA minimum

m—ssivity of the skullX @HA l—rgeY @IA sm—llF width in its middle p—rtF

PPF ‚el—tive dimensions of ™—nine to the size of the skullX @HA „he —n—lysis of the pho™id t—x— used RV unweighted ™h—r—™ters

l—rgeY @IA sm—llF —nd the footstr—p —n—lysisF pigF II shows the resulting single most

widdle wio™ene se—ls from the xorthern €—r—tethys QRQ

123456789101112131415161718192021222324 Lutra canadensis 002000000000100110110001 Devinophoca claytoni 002111100????????0011101 Cystophora cristata 111011000011021202001000 Pachyphoca ukrainica ???????????????????????? Pachyphoca chapskii ???????????????????????? Mirounga angustrirostis 112001000011222202001010 Monachus schauinslandi 112001111211112211111101 Callophoca obscura 002011000101102001010001 Pliophoca etrusca 002021010120001001110101 Pontophoca sarmatica ??????????????1???100?00 Phoca vitulina 000101121010000010111111 Leptophoca lenis 000101011111100000111101 Erignathus barbatus 010101121011111200011111

25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 Lutra canadensis 001010100101012000011011 Devinophoca claytoni 000????????????????????? Cystophora cristata 001101211001002101200111 Pachyphoca ukrainica ???102022010000001001010 Pachyphoca chapskii ????0?????1?100001?01??0 Mirounga angustrirostis 001001222110112110201211 Monachus schauinslandi 101111001111110111010111 Callophoca obscura 102110111201000010111001 Pliophoca etrusca 101110102100000011100001 Pontophoca sarmatica 0??011110001000000000110 Phoca vitulina 012021000002101011011101 Leptophoca lenis 000020000111001001001100

Erignathus barbatus 100021222011101011211111

„—˜F IH—D ˜ E w—trix for ™h—r—™terEst—te d—t— for €ho™id—e t—x— —nd outgroup —n—lyzedF

p—rsimonious ‡—gner tree gener—ted ˜y ‡in™l—d— with length of ISU humer—l tro™hle—r intern—l ™rest rising w—veElike over

stepsD gonsisten™y sndex of HFRQD —nd ‚etention sndex of HFRTF

the ™oronoid foss—D —nd the dist—l —nd proxim—l epiE

„he nodes of the ™l—dogr—m shown in pigF II —re supported ˜y

physes of the humerus —lmost equ—l in widthF the following ™h—r—™ter tr—nsform—tionsX

his™ussion of sexu—l dimorphism in ‚e™ent €hoE xode I @p—mily €ho™id—eAX T@IAF „his syn—pomorphi™ group

with —n —n™estr—l or primitive ™h—r—™ter @well developed jugul—r proE ™id—e h—s ˜een ˜—sed m—inly on ™r—ni—l m—teri—lD with

™essA is tre—ted —s plesiomorphi™ for the p—mily €ho™id—eF elsoD ™h—rE

very little —ttention to post™r—ni—l elementsF sn ™ontr—stD

—™terQQ@IDPA is homopl—sious in €—™hypho™— ukr—ini™— @whi™h ˜elongs

the fossil rem—ins of true se—ls usu—lly ™onsist of isol—ted

to the ƒu˜f—mily gystophorin—eAD €liopho™— etrus™— @whi™h ˜elongs to

lim˜ ˜onesD m—inly the humerus —nd femurF het—iled the ƒu˜f—mily won—™hin—eAD —nd irign—thus ˜—r˜—tus @whi™h ˜elongs

to the ƒu˜f—mily €ho™in—eAF studies of sexu—l dimorphism ™overing ˜oth ™r—ni—l

xode P @ƒu˜f—mily hevinopho™in—eAX R@IAF „he —nterior p—l—t—l

—nd post™r—ni—l elements in ˜oth fossil —nd ‚e™ent speE for—min— —re f—intly m—rkedF

™ies h—ve only ˜een performed on mem˜ers of the ƒu˜E

xode Q @ƒu˜f—mily €ho™in—eAX PW@PAF rumer—l tro™hle—r ™rest is

f—mily €ho™in—e @uoretsky IWVU—D PHHIAF „hereforeD this not sep—r—ted from ™oronoid foss— ˜y — distin™t lipF

xode R @ƒu˜f—mily won—™hin—eAX Q@PAY IV@IAY QI@IAF w—xill— new fossil m—teri—l provided — unique opportunity to

forms — long ™on™—vityY num˜er of in™isors redu™edY humer—l lesser

study the sexu—l dimorphism of lim˜ ˜ones —nd m—ndE tu˜er™le extends —long the ˜one9s —xisF

i˜les of the ƒu˜f—mily gystophorin—eF

xode S @ƒu˜f—mily gystophorin—eAX Q@IAY IV@PAF w—xill— forms —

ƒexu—l dimorphism in post™r—ni—l ˜ones —nd short ™on™—vityY num˜er of in™isors further redu™edY humer—l tro™hle—r

™rest r—ised w—veElike over ™oronoid foss—F elsoD ™h—r—™ter QP@PA is m—ndi˜les in living mem˜ers of the ƒu˜f—mily gystoE

simil—rin wiroung— —ngustirostris —nd €—™hypho™— ukr—ini™—F

phorin—e is more o˜vious th—n in other true se—lsF „he

qu—ntity @SV individu—l ˜ones of the post™r—ni—l skeleE

tonA —nd v—riety of the new fossil post™r—ni—l ˜ones

his™ussion —nd gon™lusions

reve—led two size ™l—sses of spe™imens th—t did not fit

known p—tterns of ontogeneti™ or sexu—l v—ri—tionF sn

„he new genus with two new spe™ies @€—™hypho™—

—dditionD most elements of the post™r—ni—l skeleton for ukr—ini™— —nd €F ™h—pskiiA of extin™t fossil true se—ls

˜oth new spe™ies reve—l sever—l very primitive ™h—r—™E

˜elongs to the p—mily €ho™id—e —nd the ƒu˜f—mily gyE

tersD su™h —sX presen™e of femor—l lesser tro™h—nter —nd stophorin—eF „hese spe™ies —re known only from the

visi˜le intertro™h—nteri™ lineY — deepD long foss— for oriE widdle wio™eneD widdle ƒ—rm—ti—n @IPFQ ± IIFP w—A

gin of the gluteus medius mus™le on the wing of the deposits of southern kr—ineF „he new wio™ene genus

iliumY wellEdeveloped pe™tine—l tu˜erosityY —nd — l—rgeD €—™hypho™— sh—res numerous ™h—r—™ters with spe™ies of

deep —™et—˜ulumF gystophor— —nd wiroung—D su™h —sX the middle of the

QRR uoretsky sFeF 8 ‚—hm—t ƒFtF

foth new spe™ies show — mos—i™ of these primiE geogr—phi™ distri˜utions to help ™l—rify phylogeneti™ reE

tive ™h—r—™tersF por ex—mpleD the sm—ller spe™ies @€—™hyE l—tionships within the ƒu˜f—mily gystophorin—e @espeE

pho™— ukr—ini™—A h—s — ˜etterEdeveloped lesser tro™h—nE ™i—lly ™omp—risons with the ™ontroversi—l genus wiE

terof the femurth—n the l—rgerspe™ies@ €F ™h—pskiiAD roung—AF „he fossil re™ord of gystophorin—e from the

while the l—rger h—s — thi™ker —nd ˜igger intertro™h—nE xorthern fl—™k ƒe— region @€—r—tethy—n f—sinA is in

teri™ ™restF „he l—rger @€F ™h—pskiiA h—s innomin—te ˜ones —greement with the e—rlier hypothesis th—t the origin

with — deepD ™oni™—l —™et—˜ulumD —nd the m—rgins of the of €ho™id—e w—s in the etl—nti™ y™e—n in™luding „ethys

—™et—˜ul—r foss— —re r—ised high —˜ove the pl—ne surf—™e @uoretsky 8 f—rnes PHHTAF hespite the in™omplete fossil

of the ˜oneF sn ™ontr—stD the sm—ller spe™ies h—s — pu˜is eviden™e for —n™ient dispers—l p—tternsD our hypothesisD

with — ˜igD wellEdeveloped ridge for —tt—™hment of the ˜—sed on this new m—teri—lD indi™—tes th—t gystophoriE

o˜tur—tor mus™les @whi™h ™—use outw—rd rot—tion of the n—e origin—ted in the €—r—tethy—n f—sinD pro˜—˜ly miE

hip jointAD — thi™kD wide —nd ro˜ust is™hi—l spine for gr—ted westw—rd —˜out IIFP million ye—rs —goD —nd su˜E

—tt—™hment of the ˜i™eps femoris mus™le @—n extensor sequently divided into the two modern gener— @gystoE

of the hip jointAD —nd — deep foss— on the medi—l —spe™t phor— —nd wiroung—AF purtherD wiroung— itself divided

of the ilium for—tt—™hment of the gluteus medius mus™le into the northern —nd southern eleph—nt se—lsF xorthern

@—lso —n extensorof the hip jointAF eleph—nt se—ls @wiroung— —ngustirostrisA spre—d through

„he presen™e of these —n—tomi™—l tr—itsD —nd speE the xorth etl—nti™ y™e—n with gystophor— ™rist—t— into

™ifi™—lly the wellEdeveloped lessertro™h—nterof the feE the er™ti™ regionF ƒouthern eleph—nt se—ls @wiroung—

murD implies th—t the sm—ller €—™hypho™— ukr—ini™— w—s leonin—A dispersed into the ent—r™ti™ region vi— the

more —d—pted to terrestri—l lo™omotion @more primitiveA ƒouth etl—nti™ y™e—nF wF —ngustirostris migr—ted

th—n its l—rger rel—tiveF sn —dditionD ˜oth new spe™ies —re through the fering ƒe— —nd l—ter on ˜e™—me the xorth

more primitive —nd ˜etter —d—pted for terrestri—l lo™oE €—™ifi™ eleph—nt se—lD while gF ™rist—t— ˜e™—me the

motion th—n —ny living represent—tives of the ƒu˜f—mily hooded se—ls of the er™ti™ gir™le —nd xorthern €—™ifi™F

gystophorin—eF xot—˜lyD the sexu—l dimorphism supE „hese new findings help expl—in the originD dispers—lD

posed in the two new spe™ies of €—™hypho™— is more —nd phylogeneti™ rel—tions —mong the sever—l su˜f—miE

o˜vious th—n the simil—r ™h—r—™ters in modern gystoE lies of the p—mily €ho™id—eF

phorin—e @see „—˜s PD TAF yurphylogeneti™ —n—lysisD ˜—sed on ™r—ni—l—nd

„his new m—teri—l of fossil se—ls from the widdle post™r—ni—l ™h—r—™ters of fossil —nd ‚e™ent se—lsD supE

wio™ene @ƒ—rm—ti—nA deposits of the kr—ine —llows ports — monophyleti™ p—mily €ho™id—e th—t in™ludes

emended di—gnosesD redes™riptionsD —nd —ssessment of the su˜f—milies hevinopho™in—eD €ho™in—eD won—™hiE

pigF II E €ostul—ted phylogeneti™ reE

l—tionships —mong sele™ted

‚e™ent —nd fossil t—x— of

the p—mily €ho™id—eD inE

™luding €—™hypho™— ukr—iE

ni™— —nd €F ™h—pskiiD new

gener— —nd new spe™iesD

˜—sed on RV ™h—r—™ters exE

pl—ined in the textF „his is

the resulting single most

p—rsimonious tree with ISU

stepsD gonsisten™y sndex

HFRQD ‚etention sndex HFRTD

o˜t—ined using the ‡in™l—E d— option of rennigVTF

widdle wio™ene se—ls from the xorthern €—r—tethys QRS pigF IP E ghronologi™ —nd str—tigr—phi™ ™orrel—tion ™h—rt of extin™t —nd living €ho™id—eF fr—n™hing points —re not ne™ess—rily ™orrel—ted with the time s™—leF

QRT uoretsky sFeF 8 ‚—hm—t ƒFtF

spe™ies r—ise some questions —˜out the —ssignment of n—eD —nd gystophorin—eF sf pho™ids —re indeed monoE

phyleti™D there ™—nnot ˜e du—l origins —nd dispers—ls other se—l m—teri—l from the €—r—tethys to €ho™— ˜esE

˜etween won—™hin—e @xorth etl—nti™ ™o—stline of s—r—˜i™—F ƒin™e ƒimiones™u @IWPSAD —ddition—l ™olle™E

xorth emeri™—A —nd €ho™in—e @€—r—tethysA —s suggested tions from the gentr—l —nd i—stern €—r—tethys h—ve

˜y hemeÂre etF —lF @PHHQAF eddition—llyD these new findE ˜een —™™umul—ted in the xwxr E€D uievD kr—ineD

È ings —ppe—rto ™ontr—di™tthe hypothesis of e rn—son etF th—t likely pert—ins to this spe™iesF „his —dded m—teri—l

—lF @PHHTAD where —ll se—ls origin—ted on the xorth h—s never ˜een des™ri˜ed or illustr—tedD with no one

etl—nti™ ™o—st of xorth emeri™—F xumerous fossil se—l even visiting the lo™—lities or rese—r™hing their str—tiE

rem—ins @—nd their geologi™—l —gesA in the €—r—tethy—nG gr—phyF

wediterr—ne—n regionsD ™olle™ted over the l—st three deE ‚esults of extensive investig—tions on these fossil

™—desD do not support either hypothesis mentioned rem—ins provide — found—tion for revision of the groupD

—˜oveF —nd ™l—rifi™—tion of the system—ti™s of its mem˜ersF

woreoverD the phylogeneti™ —n—lysis suggests th—t woreoverD these results h—ve provided the ˜—sis for reE

se—ls with IH in™isors @€ho™in—eA —re more primitive vision of the generi™ —nd spe™ifi™ di—gnoses of true se—ls

th—n those with V @won—™hin—eAD whi™h —re more primiE —nd suggest — fr—mework for the unified study of ™r—ni—l

tive th—n those with T @gystophorin—eAF elsoD it further —nd post™r—ni—l elementsD in™luding the full r—nge of

suggests th—t the ƒu˜f—mily gystophorin—e in™ludes ontogeneti™ —nd sexu—l morphologi™—l v—ri—tionsF „he

hooded @gystophor—A —nd eleph—nt @wiroung—A se—lsD methods developed in these studies ™—n ˜e —pplied

—s well —s the two new spe™ies of the widdle ƒ—rm—ti—n

equ—lly well to modern —nd fossil represent—tives of

––p—™hyosteos™leroti™99 se—ls @€—™hypho™—A @pigF IIAF

the ƒu˜f—mily gystophorin—e for des™ription —nd morE

ƒome wio™ene se—l post™r—ni—l elementsD in™luding

phometri™ tre—tmentF

ri˜sD exhi˜it thi™k —nd swollen @p—™hyosteos™leroti™A

huring our review of se—ls over—llD one tree w—s

˜ones th—t ™—n ˜e mist—ken forthose of sireni—nssu™h

ere™ted des™ri˜ing the rel—tionships of —ll ™urrently

—s w—n—tus m—eoti™us @uoretsky PHHIAF €—leoe™ologiE

known extin™t —nd living spe™iesD —s well —s the two

™—llyD it is possi˜le th—t hypers—line ™losed ˜—sins develE

spe™ies des™ri˜ed in this p—per @€—™hypho™— ukr—ini™—

oped —s the gentr—l —nd i—stern €—r—tethys ƒe— gr—du—lly

—nd €F ™h—pskiiAF „his tree w—s ™om˜ined with — ™h—rt

driedD —llowing p—™hyosteos™leroti™ se—ls —nd sireni—ns

of str—tigr—phi™ —nd ™hronologi™ ™orrel—tions @pigF IPA

to evolve independently in the s—me time periodsF pun™E

th—t —lso in™ludes multiple spe™ies th—t —re not dis™ussed

tion—llyD the in™re—sed skelet—l m—ss would ˜e — likely

in this p—per —nd sever—l undes™ri˜ed new finds th—t will

—d—pt—tion for redu™ing ˜uoy—n™y —nd to rem—in su˜E

˜e pu˜lished soonF

merged more e—sily ˜y h—ving high ˜one density @™orreE

l—ting to feeding h—˜itsAF st m—y ˜e re—soned th—t sever—l e™knowledgementsF ‡e wish to express our gr—titude to hrF

idwin qill—nd from the hep—rtment of en—tomyD gollege of wedi™ineD spe™ies evolved p—™hyosteos™leroti™ ˜ones for sh—llow

row—rd niversityD ‡—shington hgD ƒe for ™riti™—l review —nd very diving —nd slow swimming speedsF €—™hyosteos™lerosis

useful editori—l rem—rksY hrF ‰ury eF ƒemenovD for support of this

—mong fossil se—ls is — rel—tively new dis™overy —nd is

studyD ex™—v—tionD repli™—tionD —nd some photogr—phs of this m—teri—lY

h—rdly rem—rked —t —ll in liter—tureF „he det—iled study of

—nd hrF hmitry sv—noffD for re—ding —n e—rlier version of this m—nuE

p—™hyosteos™leroti™ ˜ones of fossil —nd ‚e™ent se—ls is — s™ript —nd for strengthening it with very helpful suggestions @˜oth of

pl—nned future proje™tF the l—tter —re from the hep—rtment of €—leontologyD „he x—tion—l wuE

seum of x—tur—l ristoryD x—tion—l e™—demy of ƒ™ien™eD uievD kE st is possi˜le th—t — few p—™hyosteos™leroti™

r—ineAF elsoD we w—nt to th—nk the reviewers of this p—per @hrF hF€F

˜ones des™ri˜ed ˜y ƒimiones™u @IWPSA —s €ho™— ˜ess—E

homningD hrF qF fi—nu™™iD —nd enonymousA for —ssisting in gener—ting

r—˜i™— from the i—rlyEwiddle ƒ—rm—ti—n of fess—r—˜i—

— ˜etterm—nus™riptwith theirtimeD suggestionsD —nd intelligent quesE

@modern kr—ineEwold—vi— ˜orderA —lso ˜elong to the

tionsF wost of the fin—n™i—l support for this proje™t w—s provided ˜y the

ƒu˜f—mily gystophorin—eF roweverD the l—™k of —ddiE ‚emington —nd w—rguerite uellogg pund of the ƒmithsoni—n snstituE

tion—l m—teri—ls —nd in™omplete des™ription of the type tionF

widdle wio™ene se—ls from the xorthern €—r—tethys QRU

‚ipi‚ixgiƒ

ellen tFeF @IVVHA E ristory of xorth emeri™—n €innipedsF e letzkyD „FpFD pishF ‚esF fo—rd of g—nD „r—nslF ƒerD IIRD

monogr—ph of w—lrusesD se— lionsD se— ˜e—rs —nd se—ls SU ppFD IWSUAF

of xorthern emeri™—F FƒF qeolF qeogrF ƒurvF „erritF gh—pskii uFuF @IWTIA E gurrent st—tus —nd pro˜lems in the

wis™F €u˜FD IPX IEUVSF system—ti™s of pinnipedsF „rudy ƒoveshF ikhtiolF komF

elekseev eFuF @IWPRA E ƒe—ls in the ƒ—rm—ti—n deposits of ek—dF x—uk ƒƒƒ‚D IPX IQVEIRWF

ƒouthern ‚ussi—F tF x—u™hFEsssleF k—fF v ydesseDIX gh—pskii uFuF @IWTUA E worphologi™—lEt—xonomi™—l n—ture

PTEQR ‘in ‚ussi—n“F of the p—getod form of the fering ƒe— l—rgh— se—lF snX

en˜inderiFwF @IWVHA E u—ryology—nd ivolution of the ‚ese—r™h on m—rine m—mm—ls E „rudy €oli—rF x—u™hFE

€innipedsF x—uk—D RSP ppF ‘in ‚ussi—n“F sssleF snstF ‚y˜F uhozF yke—noF @€sx‚yAF wurm—nskD

endrusov xFsF @IVWQA E qeote™toni™s of the uer™h €eninsuE PIX IRUEIUU ‘in ‚ussi—n“F

l—F w—tF qeolF ‚ussFD ITX WIEIIQ ‘in ‚ussi—n“F gh—pskii uFuF @IWUIA E ƒystem—ti™ r—nk —nd su˜generi™ difE

entoniuk eFeF @IWUWA E gomp—r—tive morphology of the ferenti—tion of se—ls @of the su˜f—mily won—™hin—eAF

—xi—l skeleton of the €innipedsF evtoreF dissertF k—ndF „rudy etl—nF x—u™hFEsssleF snstF ‚y˜F uhozF yke—no

˜iologF x—ukX IEPR ‘in ‚ussi—n“F @e„vex„xs‚yAF u—liningr—d QWX QHSEQIT ‘in ‚usE

È ern—son FD qull˜erg eFD t—nke eFD uull˜erg wFD vehm—n si—n“ @inglish tr—nsl—tionD pishF ‚esF fo—rd of g—nFD

xFD €etrov iFeF 8 †—inol— ‚F @PHHTA E €inniped €hyE „r—nsF ƒerD QIVSD IWURAF È È È

logeny —nd — new rypothesis fortheirorigin—nd gh—pskii uFuF @IWURA E sn defense of ™l—ssi™—l t—xonomy of

dispers—lF wole™F €hylF ivolD RIX QRSEQSRF the se—ls of the f—mily €ho™id—eF „rudy oolF snstF

fernorD ‚FvFD „o˜ien rF 8 ‡ood˜urne wFyF @IWWHA E €—tE e™—dF ƒ™iF ƒƒ‚F SQX PVPEQQR ‘in ‚ussi—n“F

terns of yld ‡orld ripp—rionine ivolution—ry hiE gozzoul wFeF @PHHIA E e ––northern99 se—l from the wio™ene

versifi™—tionF snX iF vinds—yD †F p—hl˜us™h 8 €F wein of ergentin—X impli™—tions of pho™id phylogeny —nd

@idsA E xe„y edv—n™ed ‚ese—r™h ‡orkshopD ˜iogeogr—phyF tF †ertF €—leontolFD PIX RISERPIF

ƒ™hloss ‚eisens˜ergD qerm—nyF iurF xeogF w—mF hemeÂre „FeFD fert— eF 8 ed—m €FtF @PHHQA E €innipediE

ghronF €lenumD xew ‰orkD ppF PTQEQIWF morph evolution—ry ˜iogeogr—phyF snX plynn vFtF

fert— eF 8 ‡yss eF‚F @IWWRA E €inniped €hylogenyF snX eF @idFA E †erte˜r—te possils —nd their gontextD gontriE

fert— 8 „FeF hemeÂre @idsA E gontri˜utions in w—rine ˜utions in ronorof ‚i™h—rdrF „edforF D fullF emerF

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fert— eF 8F ƒumi™h tFvF @IWWWA E w—rine w—mm—lsX ivoluE lo™—tions of the „erti—ry verte˜r—tes of kr—ini—n

tion—ry fiologyF e™—demi™ €ressD ƒ—n hiegoF ƒƒ‚F x—uk—D ISW ppF ‘in ‚ussi—n“F

fi—nu™™i qFD q—tt wFD g—t—nz—riti ‚FD ƒor˜i ƒFD fon—vi— gF ii™hw—ld iF @IVSHA E veth—e— ‚ossi™—F PHWEPIVD plF IQD etl—sF

qFD gurmi ‚F 8 †—rol— eF @PHIIA E ƒystem—ti™sD ˜ioE idu—rd €r—tzD ƒtF €eters˜urg ‘in ‚ussi—n“F

str—tigr—phy —nd evolution—ry p—ttern of the yligoE ii™hw—ld i @IVSQA E veth—e— ‚ossi™— ou €—leontologie de l—

wio™ene m—rine m—mm—ls from the w—ltese ssl—ndsF ‚ussieF QX IESQQF etl—sD IQ plsF ƒ™hweizer˜—rtD ƒtuttE

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finind—Eimonds yF‚F€F 8 ‚ussell eF€F @IWWTA E e morphoE inglish eF‡F @IWUUA E ƒtru™tur—l ™orrel—tes of forelim˜ fun™E

logi™—l perspe™tive on the phylogeneti™ rel—tionships tion in furse—ls —nd se— lionsF tF worphFD ISIX QPSEQSPF

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furns tFtF 8 p—y pFrF @IWUHA E gomp—r—tive morphology of n—tF felgFD PHX IERF

the skull of the ‚i˜˜on se—lD ristriopho™— f—s™i—t—D pri—nt wF @IWRUA E ‚e™her™hes sur le feÂmurdes €ho™id—eF

with rem—rks on system—ti™s of €ho™id—eF tF oolF fullF wusee royF ristF n—tF felgFD PQX IESIF Â

@vondonAD ITIX QTQEQWRF pulton „FvF 8 ƒtroke™k gF @PHIHA E wultiple m—rkers —nd

g—rr ƒFwF 8 €erry iFeF @IWWRA E sntr—E —nd interf—mili—l multiple individu—ls refine true se—l phylogeny —nd

system—ti™ rel—tionships of pho™id se—ls —s indi™—ted ˜ring mole™ules —nd morphology ˜—™k in lineF €ro™F

PUUX IHTSEIHUHF ˜y mito™hondri—l hxe sequen™esF wole™ul—r qeneE ‚oyF ƒo™F fF

hF†F @IWVPA E ƒexu—l dimorphism in stru™ture of q—djiev ti™s of w—rine w—mm—lsF ƒo™F w—rF w—mm—lFD ƒpF €u˜F

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vodnF zhivotF uur—Eer—xinF nizmenF w—lF g—u™FDTX gh—pskii uFuF @IWSPA E „he —ge —nd sexu—l ™h—nges in the

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qill „F xF @IVTTA E €rodrome of — wonogr—ph of the €inniE di—gnosis of some pinnipedsF szvestF snstF estestvenF

pedesF €ro™F issex snstFD SX QEIQF D PSX UVEWT ‘in ‚ussi—n“F x—uk

uFuF @IWSSA E en —ttempt —t revision of the sysE qins˜urg vF 8 t—nvier €F @IWUSA E ves m—mmifeÁres m—rins gh—pskii

tem—ti™s —nd di—gnosti™s of se—ls of the su˜f—mily des f—luns de l— „our—ine et de l9enjouX f—uneD giseE

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m—rine m—mm—ls of ƒ—rm—ti—n deposit —nd —djoining l— pr—n™eF xFƒFD PIXITWEIUVF

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uoretsky @auoretsk—y—A sFeF @IWVTA E wodern st—tus of sysE qr—y tFiF @IVPUA E w—mm—li— ‘volF SD PWIpF“ snX qriffith iF

tem—ti™s of the wio™ene se—ls of the p—mily €ho™id—e @idFA E „he —nim—l kingdom —rr—nged in ™onformity

of the xorthern fl—™k ƒe— €rovin™eF e˜strF sˆ ellE with its org—niz—tionD ˜y the f—ron guvierF qf ‡hitE

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in the €ho™in—e systemF €r—eprintGek—dF x—uk kr—F

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uoretsky sFeF 8 ‚—y gFiF @IWWRA E gryptopho™—D new genus

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